A Revision of New Caledonian Gossia N. Snow & Guymer (Myrtaceae)

adansonia 2020  42  7 Directeur de la publication : Bruno David Président du Muséum national d’Histoire naturelle

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Comité scientifique / Scientific board : P. Baas (Nationaal Herbarium Nederland, Wageningen) F. Blasco (CNRS, Toulouse) M. W. Callmander (Conservatoire et Jardin botaniques de la Ville de Genève) J. A. Doyle (University of California, Davis) P. K. Endress (Institute of Systematic Botany, Zürich) P. Feldmann (Cirad, Montpellier) L. Gautier (Conservatoire et Jardins botaniques de la Ville de Genève) F. Ghahremaninejad (Kharazmi University, Téhéran) K. Iwatsuki (Museum of Nature and Human Activities, Hyogo) K. Kubitzki (Institut für Allgemeine Botanik, Hamburg) J.-Y. Lesouef (Conservatoire botanique de Brest) P. Morat (Muséum national d’Histoire naturelle, Paris) J. Munzinger (Institut de Recherche pour le Développement, Montpellier) S. E. Rakotoarisoa (Millenium Seed Bank, Royal Botanic Gardens Kew, Madagascar Conservation Centre, Antananarivo) É. A. Rakotobe (Centre d’Applications des Recherches pharmaceutiques, Antananarivo) P. H. Raven (Missouri Botanical Garden, St. Louis) G. Tohmé (Conseil national de la Recherche scientifique Liban, Beyrouth) J. G. West (Australian National Herbarium, Canberra) J. R. Wood (Oxford)

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© Publications scientifiques du Muséum national d’Histoire naturelle, Paris, 2020 ISSN (imprimé / print) : 1280-8571/ ISSN (électronique / electronic) : 1639-4798 A revision of New Caledonian Gossia N. Snow & Guymer (Myrtaceae)

Neil SNOW Department of Biology, T.M. Sperry Herbarium 1701 S. Broadway St., Pittsburg State University Pittsburg, KS 66762 (United States) [email protected]

Submitted on 28 June 2018 | accepted on 21 September 2019 | published on 27 April 2020

Snow N. 2020. — A revision of New Caledonian Gossia N. Snow & Guymer (Myrtaceae). Adansonia, sér. 3, 42 (7): 131-177. https://doi.org/10.5252/adansonia2020v42a7. http://adansonia.com/42/7

ABSTRACT Gossia N. Snow & Guymer from New Caledonia as revised here comprises 30 taxa. Nine new species are proposed, including: Gossia angustifolia sp. nov., G. bourailensis N. Snow, sp. nov., G. conduplicata sp. nov., G. kaalaensis sp. nov., G. katepahiensis sp. nov., G. mandjeliaensis sp. nov., G. ngaensis sp. nov., G. ouazangouensis, and G. ramiflora sp. nov. Ten new subspecies are proposed: two in Gossia aphthosa (Vieill. ex Brongn. & Gris) (G. a. subsp. longipedunculata N. Snow & Munzinger, subsp. nov. and subsp. austro-orientalis N. Snow & K. Gandhi, subsp. nov.), and eight in Gossia clusioides (Brongn. & Gris) N. Snow (G. c. subsp. avanguiensis subsp. nov., G. c. subsp. bleuensis subsp. nov., G. c. subsp. callmanderiana subsp. nov., G. c. subsp. maoyana subsp. nov., G. c. subsp. ploumensis N. Snow, comb. et stat. nov., G. c. subsp. rembaiensis subsp. nov., G. c. subsp. taomensis subsp. nov., and G. c. subsp. tiebaghiensis subsp. nov. The new combination Gossia conspicua (Vieill. ex Guillaumin) N. Snow, comb. nov. is made; a second-stage lectotype and epitype are selected for Gossia diversifolia (basionym Eugenia diversifolia Brongn. & Gris); a lectotype is selected for Eugenia heckelii Pancher & Sebert, which here is reduced to synonymy under G. vieillardii (Brongn. & Gris) N. Snow; and a lectotype is selected for Eugenia cataractarum Guillaumin, which here is reduced to synonymy under KEY WORDS G. kuakuensis (Baker f.) N. Snow. Myrtus pulchrefolius Guillaumin is reduced to synonymy under New Caledonia, Myrtaceae, Gossia alaternoides. The Australian speciesGossia byrnesii N. Snow & Guymer is synonymized under conservation, Eugenia reinwardtiana DC. The treatment provides a key to and descriptions of all known New Cal- lectotypification, edonian taxa and includes comments on distributions, distribution maps, ecology and phenology. new combinations, new subspecies, Illustrations are provided for nearly all taxa and digital images of living material are presented for new species. several. With this revision, Gossia now comprises 45 species and ten subspecies.

ADANSONIA, sér. 3 • 2020 • 42 (7) © Publications scientifiques du Muséum national d’Histoire naturelle, Paris. www.adansonia.com 131 Snow N.

RÉSUMÉ Révision du genre néo-calédonien Gossia N. Snow & Guymer (Myrtaceae). Le genre Gossia N. Snow & Guymer de Nouvelle Calédonie, révisé ici, comprend 30 taxons. Neuf espèces nouvelles sont proposées, parmi lesquelles : Gossia angustifolia sp. nov., G. bourailensis N. Snow, sp. nov., G. conduplicata sp. nov., G. kaalaensis sp. nov., G. katepahiensis sp. nov., G. mandjeliaensis sp. nov., G. ngaensis sp. nov., G. ouazangouensis, et G. ramiflora sp. nov. Dix nouvelles sous-espèces sont proposées : deux issues de Gossia aphthosa (Vieill. ex Brongn. & Gris) (G. a. subsp. longipedunculata N. Snow & Munzinger, subsp. nov. et subsp. austro-orientalis N. Snow & K. Gandhi, subsp. nov.), et huit de Gossia clusioides (Brongn. & Gris) N. Snow (G. c. subsp. avanguiensis subsp. nov., G. c. subsp. bleuensis subsp. nov., G. c. subsp. callmanderiana subsp. nov., G. c. subsp. maoyana subsp. nov., G. c. subsp. ploumensis N. Snow, comb. et stat. nov., G. c. subsp. rembaiensis subsp. nov., G. c. subsp. taomensis subsp. nov., and G. c. subsp. tiebaghiensis subsp. nov. Une nouvelle combinaison, Gossia conspicua (Vieill. ex Guillaumin) N. Snow, comb. nov., est proposée ; un lectotype de seconde étape et un épitype sont sélectionnés pour Gossia diversifolia (basionyme : Eugenia diversifolia Brongn. & Gris); un lectotype est désigné pour Eugenia heckelii Pancher & Sebert, qui est ici mis en synonymie avec G. vieillardii (Brongn. & Gris) N. Snow; enfin, un lectotype est désigné pourEugenia cataractarum Guillaumin, qui est mis ici en MOTS CLÉS synonymie avec G. kuakuensis (Baker f.) N. Snow. Myrtus pulchrefolius Guillaumin est mis en synonymie Nouvelle Calédonie, Myrtaceae, avec Gossia alaternoides. L’espèce australienne Gossia byrnesii N. Snow & Guymer est mise en synonymie conservation, avec Eugenia reinwardtiana DC. Une clé d’identification, ainsi que les descriptions de tous les taxons lectotypification, connus de Nouvelle Calédonie sont proposées. Enfin, des commentaires sur la distribution, l’écologie combinaisons nouvelles, sous-espèces nouvelles, et la phénologie, ainsi que des cartes de répartition des taxons cités sont fournis, qui sont également, espèces nouvelles. presque tous, figurés ici. Avec cette révision, Gossia contient maintenant 45 espèces et dix sous-espèces.

INTRODUCTION ful assessments require well-resolved and current systematic treatments, knowledge of the sizes of populations and their Gossia N. Snow & Guymer (Myrtaceae) ranges from the east- distributions and threats, have well as trends of the sizes of ern parts of New Guinea and Australia to New Caledonia and populations and knowledge about which populations occur the Solomon Islands, with most taxa occurring in Australia in protected areas (IUCN 2012). The purpose of this revision or New Caledonia (Snow et al. 2003; Snow 2005, 2006; is to propose numerous new taxa, provide a key to the New Snow & Wilson 2010; Snow & Veldkamp 2010). Lenwebbia Caledonian taxa and detailed descriptions and synonymy for N. Snow & Guymer and Gossia recently were segregated from all taxa, drawings of most taxa, digital images of living material Austromyrtus (Nied.) Burret, which formerly had a broader for some taxa, a hotlinked representative herbarium specimen circumscription; Austromyrtus and Lenwebbia as revised are for each taxon, and cite all confirmed specimens. Detailed endemic to Australia (Snow et al. 2003). IUCN (2012) recommendations are not included, but broad New Caledonia was one of the original “hotspot” areas, recog- statements are given that include relative abundance, distribu- nized for its exceptional levels of overall diversity and high levels of tion, and species needing more study for detailed assessments. plant endemism (e.g., Myers 1988). More recently, authors have proposed the concept of hotspots within hotspots (Cañadas et al. 2014) and micro- and nano-hotspots (Fenu et al. 2010). Gâteblé MATERIAL AND METHODS et al. (2018) documented the heuristic value of these concepts in their overview of taxonomic novelties in New Caledonia since The author has studied New Caledonian Gossia intermittently 2000 and concomitant description of eight new species from Île since 1998, including fieldwork on Grande Terre in 2003, ap- Art in the Belep Archipelago, where unfortunately vegetation proximately ten weeks of herbarium studies at P (2003, 2005 destruction also is rampant. Whilst numerical tallies shift as taxa and 2015), numerous shorter-duration visits to MO, and study are discovered and existing names are reduced to synonymy, the of specimens on-site while in residence at GREE, BISH, and Myrtaceae without question are at or near the top of the vascu- KSP (abbreviations following Thiers 2019). Concepts of spe- lar plant families regarding overall diversity of species in New cies and infraspecific taxa follow the basic principles outlined in Caledonia (e.g., Morat et al. 2012). With further study, some Snow (1997) and their later modification to reflect the General of the taxa proposed here likely will qualify as microendemics. Lineage Concept (de Queiroz 1998) and its application using a Prior to this treatment, Gossia comprised approximately diagnosability criterion (Snow et al. 2003; Johnson & Johnson twenty-seven accepted species and two non-nominal varie- 2006; Johnson et al. 2012). Given that many plant species in ties. Botanists and conservation biologists have intensified New Caledonia have high fidelities to certain geological sub- efforts recently to provide rigorous assessments following strates, and that certain areas of the country are particularly IUCN (2012) criteria in New Caledonia, in accordance with high in levels of microendemism, additional consideration has the recent global emphasis on plant conservation. Meaning- been accorded to substrate and geographical occurrences. The

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paucity of material for some taxa is reflected by taxonomic de- Branchlets scriptions that are not always strictly in parallel. Terminology Terete, compressed, or rarely 4-angled. largely follows my recent treatments for New Caledonia (Snow et al. 2016a, b) or Beentje (2010). Measurements refer to lengths Colleters or length × width. All specimens cited have been seen unless Absent to obscure, reddish and hair-like if present. indicated with n.v.; barcodes [in square brackets] are indicated for many specimens, especially those at NOU or P; accession Leaves numbers are hyphenated (e.g., MO-5906585). Specimens are 2(-3-4) per node, pedicellate or sometimes sessile, venation cited alphabetically by senior collector in their ascending order brochidodromous, thinly to thickly coriaceous, blades flat numerically, from the North (Nord) and South (Sud) Provinces, or bullate. respectively. Geo-coordinates are given for selected specimens; those in square brackets were determined post facto by various Inflorescence workers, mostly at NOU. Geocoordinates and maps of many Axillary, terminal or ramiflorus, of monads, racemes, or pani- specimens can be seen by entering the relevant barcode of speci- cles, solitary, paired, or fascicled; extrafloral bracts absent, mens at P (see: https://science.mnhn.fr). Images of living taxa scale-like or sometimes leafy. indicated as “vel aff.” are best estimates of their identities given the lack of voucher specimens. For researchers new to plant Bracteoles taxonomy in New Caledonia, Hopkins & Bradford (2009) is 2, sometimes early caducous. worthy of review given their discussion of problems with some previous numbering systems. As applied here, the largest num- Hypanthium ber of separate taxa initially assigned to one number (including Campanulate, cupulate, obconic, or urceolate, not prolonged members of other genera) was Balansa 1490. All specimens from beyond ovary apex, smooth or sometimes ribbed or rugose; WELTU, including those used in illustrations, recently were ovary apex glabrous to hairy. transfered to WELT (B. Sneddon pers. comm. 2017). Several species are widespread and surely not of conservation concern. Flowers Others with fewer specimens should be assessed following IUCN 4-5(-8)-merous. (2012) criteria and additional fieldwork. Locules (1-)2(-3), placentation axile (or basal in G. angustifolia sp. nov.). SYSTEMATICS Stamens numerous; filaments dorsifixed or subdorsifixed. Stigma apex terete, narrow to slightly globular. Genus Gossia N. Snow & Guymer Fruit Distribution. — Gossia is widespread across Grande Terre in Baccate, green (immature) to yellow-orange or red but usually New Caledonia and occasional on adjoining islands, but so far is unreported for the Loyalty Islands; mostly in maquis or more mesic purplish-black at maturity. forests and species rarely occurring in dense or extensive populations; near sea level to 1500 m. Seeds 1-4(-12), testa hard, irregularly rounded and typically Generic description of Gossia somewhat compressed with some angular edges; embryo Plants coiled or less commonly C-shaped. Chromosome numbers Shrubs to large trees, (0.2-)0.5-18 m. unknown.

Artificial key to the New Caledonian species of Gossia N. Snow & Guymer 1. Branchlets 4-angled (check youngest material) ...... G. diversifolia (Brongn. & Gris) N. Snow — Branchlets rounded or compressed ...... 2 2. Leaf blades blades broadly linear to very narrowly elliptic, apex retuse ...... 3 — Leaf blades broader than narrowly elliptic, apex obtuse to acute (rarely retuse) ...... 5 3. Leaves 4 per node, blades broadly linear to very narrowly obovate ...... G. virotii (Guillaumin) N. Snow — Leaves 2(-3) per node, blades narrowly elliptic to broader ...... 4 4. Leaves thickly coriaceous, margins strongly revolute; internodes at branch tips mostly less than 5 mm long ...... G. conspicua (Vieill. ex Guillaumin) N. Snow, comb. nov. — Leaves coriaceous, margins flat to only slightly revolute; internodes at branch tips mostly greater than 5 mm long ...... G. alaternoides (Brongn. & Gris) N. Snow 5. Flowers all or mostly 4-merous ...... 6 — Flowers all or mostly 5-merous ...... 10

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6. Leaf blades (8.5-)12 cm or longer ...... G. aphthosa (Vieill. ex Brongn. & Gris) N. Snow (in part) — Leaf blades less than 9.5 cm ...... 7 7. Leaf blades mostly 6-9.5 cm ...... G. angustifolia N. Snow, sp. nov. — Leaf blades mostly < 6 cm ...... 8 8. Leaf base narrowly cuneate; petiole sometimes drying nearly black ...... G. nigripes (Guillaumin) N. Snow — Leaf base rounded to cordate and conduplicate; petiole drying brownish ...... 9 9. Leaf blade conduplicate only at base, surface flat to slightly undulate; apex of inflorescence rigid, not nodding ...... G. bourailensis N. Snow, sp. nov. — Leaf blade conduplicate up to half or more of its length, surface moderately to strongly undulate; apex of inflorescence sometimes nodding ...... G. conduplicata N. Snow, sp. nov. 10. Petals > 9 mm ...... 11 — Petals < 9 mm (petals unknown in G. conduplicata N. Snow, sp. nov., which probably keys here) ...... 13 11. Base of leaf cordate to rounded; petiole 1.5-2 mm ...... G. aphthosa subsp. longipedunculata N. Snow & Munzinger, subsp. nov. — Base of leaf round to cuneate, petiole 2-30 mm ...... 12 12 Calyx lobes sparsely sericeous above; petals sericeous-tomentose below (sometimes densely so); hypanthium surface smooth; leafy bracts along inflorescence axis prominent (but typically soon caducous) ...... G. clusioides (Brongn. & Gris) N. Snow — Calyx lobes sparsely ciliate only on margins; petals minutely ciliate apically but otherwise glabrous below; hypanthium surface sometimes slightly ribbed; leafy bracts along inflorescence mostly absent ...... G. colnettiana (Guillaumin) N. Snow 13. Base of leaf blade moderately to strongly conduplicate, blade surface undulate ...... G. conduplicata N. Snow, sp. nov. — Base of leaf blade more or less flat (slightly conduplicate inG. katepahiensis N. Snow, sp. nov.), surface mostly flat...... 14 14. Leaf blades mostly 10 cm or longer ...... 15 — Leaf blades mostly 10 cm or shorter ...... 16 15. Base of petiole sometimes encircled by a corky ring; base of leaf blade cordate to rounded, sometimes clasping; lowermost inflorescences often ramiflorous below leaves; bracteoles very narrowly elliptic to very narrowly obovate ...... G. aphthosa (Vieill. ex Brongn. & Gris) N. Snow (in part) — Base of petiole lacking corky outgrowths; base of leaf rounded or cuneate, never clasping; lowermost inflorescences axillary; bracteoles narrowly ovate to ovate...... G. clusioides (Brongn. & Gris) N. Snow 16. Leaf surface bullate (sometimes strongly) or margins strongly and irregularly undulate; bark of second and third year growth often dark and breaking in irregular rectangle patches ...... 17 — Leaf surface smooth or margins to flat or somewhat undulate; bark of second and third year growth mostly somewhat smooth, grayish-brown (or dark and flaking inG. kuakuensis or G. pancheri (Brongn. & Gris) N. Snow) ...... 18 17. Leaf margins more or less flat; lowermost inflorescences axillary, extrafloral bracts of inflorescence leafy; hypanthium sparsely sericeous in flower; petals sericeous to tomentose below ...... G. clusioides (Brongn. & Gris) N. Snow (in part) — Leaf margins irregularly undulate; lowermost inflorescences on naked branches below leaves, extrafloral bracts absent or scale-like; hypanthium sparsely to moderately sericeous in flower; petals ciliate...... G. ramiflora N. Snow, sp. nov. 18. Leaf apex mostly retuse (rarely obtuse) ...... 19 — Leaf apex obtuse to acute ...... 20 19. Hypanthium c. 3 mm; bracteoles and calyx lobes sparsely to densely sericeous; leaves 2 per node ...... G. pancheri (Brongn. & Gris) N. Snow — Hypanthium 2.0-2.5 mm; bracteoles and calyx lobes hairy; leaves 2 or 3 per node ...... G. alaternoides (Brongn. & Gris) N. Snow 20. Calyx lobes drying mostly whitish or cream-colored (“petaloid” of some authors) ...... 21 — Calyx lobes drying mostly greenish ...... 22

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21. Leaves mostly over 8 cm long, apex acute; adaxial leaf surface somewhat glossy, secondary veins prominently raised; margin somewhat undulate ...... G. kaalaensis N. Snow, sp. nov. — Leaves mostly shorter than 7 cm long, apex obtuse; adaxial leaf surface matte, secondary veins flush; margin slightly revolute ...... G. ouazangouensis N. Snow, sp. nov. 22. Leaf base cordate or round ...... 23 — Leaf base cuneate to narrowly cuneate ...... 27 23. Hypanthium sparsely to moderately sericeous ...... 24 — Hypanthium glabrous or very sparsely sericeous near apex ...... 25 24. Extrafloral bracts of inflorescence leafy; bracteoles c. 3.5 mm long; petals 8-13 mm ...... G. colnettiana (Guillaumin) N. Snow — Extrafloral bracts of inflorescence leafy; bracteoles scale-like, 0.5-1.4 mm long; petals (4-)5-7 mm...... G. vieillardii (Brongn. & Gris) N. Snow 25. Older branchlets with dark, rectangular flakes; extrafloral bracts of inflorescence leaf-like...... G. kuakuensis (Baker f.) N. Snow — Older branchlets smooth; extrafloral bracts of inflorescence lacking or small and scale-like...... 26 26. Petioles 1-3 mm, rounded above; leaf apex broadly acute to obtuse; ovary apex hairy ...... G. bourailensis N. Snow, sp. nov. — Petioles 2.5-4 mm, sometimes slightly sulcate; leaf apex broadly rounded or slightly retuse; ovary apex glabrous...... G. katepahiensis N. Snow, sp. nov. 27. Secondary veins on adaxial leaf (in sicco) surface prominently raised ...... 28 — Secondary veins on adaxial leaf surface (in sicco) flush or only slightly raised...... 29 28. Leaves elliptic, apex rounded (occasionally retuse); hypanthium sericeous, oil glands of uniform size; ovary apex densely tomentose ...... G. pancheri (Brongn. & Gris) N. Snow — Leaves narrowly elliptic to narrowly ovate or somewhat elliptic, apex acute; hypanthium mostly glabrous, oil glands large and small; ovary apex glabrous ...... G. kaalaensis N. Snow, sp. nov. 29. Oil glands on abaxial leaf surface absent to sparse; style villous lower ½-¾, sometimes densely so; base of fruit rounded or often prominently tapered ...... G. vieillardii (Brongn. & Gris) N. Snow — Oil glands on abaxial leaf surface dense; style sericeous to tomentose; base of fruit rounded ...... 30 30. Hypanthium sericeous ...... G. mandjeliaensis N. Snow, sp. nov. — Hypanthium glabrous ...... 31 31. Petioles reddish (drying dark brown); mature leaves mostly 5.0 cm or longer, surface flat, margin slightly recurved; Île des Pins ...... G. ngaensis N. Snow, sp. nov. — Petioles dark red (drying nearly black); mature leaves mostly 4.5 cm or shorter, blades undulate, margin flat; Massif du Panié ...... G. nigripes (Guillaumin) N. Snow

Gossia alaternoides (Brongn. & Gris) N. Snow Etymology. — Possibly from a superficial resemblance to the (Figs 1-3) Mediterranean species Rhamnus alaternus L., but its exact application here uncertain. Austrobaileya 8 (2): 180 (2010). — Myrtus alaternoides Brongn. & Gris, Bulletin de la Société botanique de France 12: 177 (1865). — Distribution, habitat and phenology. — Across much of Grande Austromyrtus alaternoides (Brongn. & Gris) Burret, Notizblatt des Terre (Fig. 3) but most common in the South Province; in maquis Botanischen Gartens und Museums zu Berlin-Dahlem 15: 504 (1941). — including over ultramafics, 100-700 m. Flowering November through Typus: New Caledonia. Grande Terre, South Prov., Montagnes de May; in fruit yearround but evidently peaking in April and May. Balade, 1855-1860, E. Vieillard 495 (holo-, P[P00602575]). Specimens examined.— New Caledonia. Grande Terre, North Gossia alaternoides (Brongn. & Gris) N. Snow var. pulchrifolius Prov., au-dessus d’Ouroué à l’embouchure du Dotio, I.1872, Bal- (Guillaumin) N. Snow, Austrobaileya 8: 180 (2010). — Myr- ansa 3398 (P[P00459516]); Goro-Nickel, relevé 37-38, 20.VI.2002, tus pulchrifolius Guillaumin, Mémoires du Muséum national Dagostini 525 (NOU); Fetscherin s.n., 1910, entre Thio et Houaïlou d’Histoire naturelle, Paris, sér. B., Botanique 8: 144 (1959); (P[P00459523]); Route de Houaïlou à Kouaoua, vers 400 m, Mémoires du Muséum national d’Histoire naturelle, Paris, sér. 7.V.1969, Jaffré 231, (BISH[fragment], NOU[NOU053232], B., Botanique 8: 289 (1962) (as M. pulchrefolius). — Typus: P[P00462828]); Boulinda, 100 m, 9.XI.1971, Jaffré 480 New Caledonia. Grande Terre, South Prov., au-dessus du (NOU[NOU028887], P[P00462829]); Massif du Boulinda, 530 m, campement de Bernier, Montagne des Sources, 3.X.1951, 12.X.1972, Jaffré 973(NOU[NOU028885], P[P00462831]); Hürlimann 3021 (holo-, P[P00751827]; iso-, A[A00255461], Au pied du Massif du Boulinda, 110 m, 10.X.1972, Jaffré 1030 NY, US, Z[Z-000050850]). (NOU[NOU028853]); Plaine des Gaiacs, 10.XII.1977, Jaffré

ADANSONIA, sér. 3 • 2020 • 42 (7) 135 Snow N.

F C

KL H J

Fig. 1. — Gossia alaternoides (Brongn. & Gris) N. Snow, showing variation in the degree of indumentum (glabrous to sericeous or tomentose) and leaf shape: A, branchlet; B, sericeous flower with bracteoles, hypanthium, calyx lobes, petals, tips of stamens, and apex of stigma; C, calyx lobes, staminal disk, points of petal attachment, and style; D, young fruit (left), mature seed (right); E, branchlet; F, leaves and flowers in bud;G , glabrous flower with detail of bracteoles and hypanthium in bud; H, branchlet; I, inflorescence; J, tomentose flower with bracteoles; K, branchlet; L, revolute leaf and young fruit. Vouchers: A-C, Jaffré 141 (NOU); D, Nothis 331 (NOU); E-G, Hürlimann 3021 (P); H-J, Jaffré 3275 (NOU); K-L, Jaffré 231 (NOU). E-G, reflect what previously was calledGossia alaternoides var. pulchrefolius. Scale bars: A, E, H, K, 2 cm; B, C, G, J, 2 mm; D, 3 mm; F, I, L, 5 mm.

136 ADANSONIA, sér. 3 • 2020 • 42 (7) Revision of New Caledonian Gossia N. Snow & Guymer

Fig. 2. — Gossia alaternoides (Brongn. & Gris) N. Snow: A, young leaves with tomentose indumentum and flower;B , Gossia diversifolia, close-up of flowers and buds, unvouchered taken in Parc Forestier de Nouméa. Vouchers: A, Callmander 1273, used with permission; B, photo: ©Benoît Henry, used with permission (http://endemia.nc).

2018 (NOU[NOU028886]); Massif du Boulinda, col de Nékoro, Kee 44240 (BRI, MO[MO-6751071], NOU[NOU028855], 11.X.1972, Jaffré 2203 (NOU[NOU028884]); Massif du Boulinda, P[P02089888] + spirits); Port Bouquet, Rivière Koum, au bout 23.II.1978, Jaffré 2367(NOU[NOU028852], P[P00462833]); de la piste longeant la rivière, berge ouest, 105 m, 24.XI.2001, Koniambo, 534 m, 26.X.2000, Jaffré 3452 (NOU[NOU028919], Munzinger 1019 (MO[MO-04771341], NOU[NOU028880]); P[P00316477]); Mt. Koniambo, vers 400 m, 5.I.1961, MacKee Dothio: le fort Américain, 17.XII.1981, Veillon 4767 (NOU, 7965 (CANB, P[P00462840], US); Boulinda, 10.XI.1977, Morat P[P00462845]); Tontouta, rive gauche du creek Wanewano entre 5705 (NOU[NOU028876], P[P00462843]); Dothio: le fort amé- 20-30 m, 26.III.1993, Veillon 7917 (NOU[NOU028920]); Baie ricain, [21°35’30.00”S, 166°08’09.99E], 17.XII.1981, Veillon 4767 de Prony, Vieillard 2618 (P[P00462847]); Entre Goro et la Grand (NOU[NOU030916], P[P00462845], WELTU). — South Prov., Lac, 29.III.1942, 230 m, Virot 680 (P[P00462851]); Riv. des Lacs Col de Vulcain, 900 m, 11.XI.1950, Baumann-Bodenheim 8120 near beginning of Route de Carenage, 22.10°S, 166.50°E, 150 m, (A, P[P00459518]); Sommet W du Mt. Humboldt, 21.IX.1951, 11.XII.1973, Webster 19208 (DAV, P[P00462852]). — Ab. loco: Baumann-Bodenheim 15430 (A, BRI, P[P00459521], US, Z); Mine Le Rat & Le Rat 768 (P[P00462835]). Pin-Pin, 21°22’57.00”S, 165°16’00.00”E, 677 m, 20.XI.2015, Callmander 1273 (G[images of living material seen], MO n.v., Representative herbarium specimen. — P00751827. NOU n.v., P n.v.); Massif du Kouakoué sur la crête, VII.1955, Chevalier 6 (NOU[NOU028915, NOU028921]); Goro-Nickel Description relevé 37-38, 20.VI.2002, Dagostini 525 (NOU[NOU028874]); Shrubs Prony, bords de la Yaté, Franc 1787 (P[P00459524], [P00459525]; ibid. loco, Franc 1787a (P[P00459526, P00462814, P00462815]); 0.5-3 m. Prony, 20.XII.1914, Franc 1901a (A, BRI, P); Hürlimann 597 (P[P00462820]); Vallée de la Fausse Yaté, 12.I.1951, Hürlimann Branchlets 669 (A, P[P00462821], RSA, US, Z); Pic 576 au-dessus de la vallée Terete, emerging densely lanate to glabrous; internodes at de Boulari, 30.I.1951, Hürlimann 799 (A, NY, P[P00462822], US, branch tips mostly > 5 mm. Z); Pic 576 au-dessus de la vallée de Boulari, 30.I.1951, Hürlimann 802 (P[P00462823]); Pente nord du Pic du Casse-Cou, 3.III.1951, Hürlimann 1016 (P[P00462825]); Route de Yaté, 10.I.1969, Leaves Jaffré 141 (NOU[NOU053233], P[P00462826, P00462827]); Stiffly coriaceous, 2-3(-4) per node; petioles 1.5-2.5 mm, Kouaoua, 7.V.1969, Jaffré 231 (P[P00462828]); Route de Yaté, slightly sulcate distally; blades 1.5-5.7 × (0.6-)1.2- 20.I.1978, Jaffré 2254(CANB n.v., L n.v., MO[MO-5813358], 2.5 cm, narrowly obovate (or rarely obovate [e.g., Vieil- NOU[NOU028883], P[P00462832]); Vallée de la Riv. Ouha-Tontou- ta, 2.XI.1995, Jaffré 3275 (BISH[fragment], NOU[NOU053039]), lard 2618]) to narrowly elliptic or elliptic, base cuneate P[P00462834]); N. Dumbéa valley between the 1st and 4th bunkers to attenuate, surface flat in center but margin typically of abandoned mine, 13.XI.1955, MacKee 3337 (P[P00462837, recurved (often strongly) throughout, apex prominently P00462838]); Plaine des Lacs: NE Grand Lac, 30.XII.1978, Mac- retuse or less often obtuse, midnerve slightly sulcate Kee 36304 (NOU[NOU028854]); Yaté, 250 m, 18.XII.1985, proximally becoming more or less flush, secondary veins MacKee 42946 (MO[MO-6751083], NOU[NOU028879], P[P02089890]); 5 km en aval de la Chute de la Rivière des Lacs, invisible above, oil glands below (when not obscured 200 m, 9.XI.1989, MacKee 43339 (BRI, NOU[NOU028875], by indumentum on branchlets) common to dense but P[P02089861] + spirits); Rivière des Lacs, 5 km en aval de la often indistinct. chute, 200 m, 5.IV.1987, MacKee 43489 (MO[MO-6751077], NOU[NOU028887], P[P02089889]); Yaté, Barrage, 150 m, Inflorescence 25.V.1987, MacKee 43555 (P[P02089859]); Rivière des Lacs, 220 m, 19.XI.1987, MacKee 43776 (P[P02089860]); Grand Lac, 2-4 cm, of monads, triads or few-flowered cymes, terminal 250 m, 29.XII.1989, MacKee 44239 (BRI, NOU[NOU028878], or (mostly) axillary, pedicels 0-6 mm; extrafloral bracts P[02089887] + spirits); Grand Lac, 250 m, 29.XII.1988, Mac- typically at base of inflorescence branches.

ADANSONIA, sér. 3 • 2020 • 42 (7) 137 Snow N.

Bracteoles Gossia angustifolia N. Snow, sp. nov. 0.6-1.2 mm (occasionally two pairs in decussate arrangement (Figs 3; 4A-C) below solitary flowers), very narrowly obovate, glabrous to densely lanate. Leaves narrowly elliptic, cuneate at base, the margin slightly sinuous, about three times longer than broad, abaxial surface glands dense, placentation basal. Hypanthium 2.0-3.0 mm, cupulate, smooth to slightly ribbed, glabrous to Typus. — New Caledonia. Grande Terre, North Prov. Entre Tao et densely lanate; ovary apex densely hairy. Ouaième: Vallée Pouai, 20°35’33.00”S, 164°49’12.00”E, 11.IX.1961, MacKee 15612 (holo-, NOU[NOU053037]; iso-, BISH[leaf fragment], MO[MO-5906585]). Calyx Lobes 5, (0.6-)1-2 mm, narrowly obovate to narrowly oblong, Paratypi. — New Caledonia. Grande Terre, Haute Diahot, Paala, 600 m, glabrous to velutinous above and below. 15.VII.1977, MacKee 33461 (BISH[fragment], NOU[NOU053245]); Col d’Amos, 20°17’60”S, 164°25’42”E, 13.X.2002, Munzinger 1489 et al. (MO[MO-5841144], NOU[NOU002733], P[P00354502]); Col Petals d’Amoss, 20°18’22”S, 164°23’48”E, 14.X.2002, Tronchet 387 (K n.v., (1.4-)2-4 × 2.5-3.5 mm, glabrous to densely hairy above and MO[MO-5906585], NOU[NOU003170], P[P00354930]); Vallée de below, white to maroon. Néouna (Goapin), entre Aoupinié NW et Forêt plate, 165°13’35.1”S, 21°8’38.3”E, 31.VII.2012, Vandrot 597 & Chambrey (MO, n.v., MPU[MPU028535], P[P02089718]). Filaments 2-3 mm, white; anther sacs (0.1-)0.3-0.7 mm, stramineous. Etymology. — In reference to the relatively narrow leaves, compared to most congenerics in New Caledonia.

Style Representative herbarium specimen. — P00354930. 4-5 mm, sometimes hairy proximally, whitish. Distribution, habitat and phenology. — Gossia angustifolia Fruits sp. nov. occurs on the edge of the Massif du Panié and at Col d’Amoss (Fig. 3), sometimes common locally; in gallery forests over micaschistes, c. 4.0-5.5 × 4 mm, globose to subcylindrical, yellow-green c. 100-600 m altitude. Flowering November (material scant); fruiting maturing dark bluish-black. July through October.

Seeds Description Tw o. Shrubs or trees 3-8 m, sometimes densely branched. Remarks The indumentum on the emerging branchlets, young leaves, Bark and floral parts varies considerably; in many specimens most Brownish and somewhat rough. or all of it falls away with age. Three or sometimes four leaves per node often occur on this generally sparsely-branched Branchlets shrub. The apex of the bracteoles in some specimens exceeds Terete to laterally compressed; internodes > 5 mm., emerging the base of the calyx lobes, but this character is inconsistent. reddish. The dried immature fruits can appear to be almost capsular and only slightly fleshy. Additional research is needed to test Leaves the distinctness of G. alaternoides, G. conspicua comb. nov., Coriaceous, 2 per node; petioles 1.2-3.5 mm, terete; blades and G. virotii. One focal area in particular for additional col- 5.0-9.0 × 1.9-3.3 cm, narrowly elliptic, base cuneate, surface lections should be near Tontouta, an area from which speci- flat, margin flat to slightly wavy, apex acute, midnerve above mens can be challenging to attribute to these three species. sulcate, secondary veins slightly raised above, oil glands of lower Previously (in ms.), I recognized up to four infraspecific surface dense but faint. Inflorescence 2-2.5 cm, of few-flowered taxa of this species, but further study only corroborated the racemes or panicles, terminal and axillary; pedicels 0.3-1.4 cm; distinctness of Gossia conspicua comb. nov., in contrast to extrafloral bracts lacking or scale-like. tentatively having recognized G. a. subsp. pulchrefolius and earlier tentative names of Gossia alaternoides vars. “tomentosa” Bracteoles and “velutinosa”. Many specimens bear annotations of those 1.5-2 mm, narrowly oblanceolate, sparsely sericeous. tentative names between 2004 and 2017. Sterile specimens of G. alaternoides can resemble Uromyrtus Hypanthium emarginata (Pancher ex Brongn. & Gris) Burret, but that 2.5-3.5 mm, campanulate or urceolate, width in flower un- species has pendulous solitary flowers. The leaves also can known, surface smooth, sparsely sericeous; ovary apex glabrous. resemble Eugenia stricta Pancher ex Brongn. & Gris, but its vegetative and reproductive parts are glabrous and the Calyx embryos are globular and lack the hardened testa charac- Lobes 5, 1.2-1.5 mm, broadly rounded, glabrous above, sparsely teristic of Gossia. sericeous below, greenish.

138 ADANSONIA, sér. 3 • 2020 • 42 (7) Revision of New Caledonian Gossia N. Snow & Guymer

165°E 166° N

Mt. Panié Ouvéa (1629 m) Island

21°S Koné 21°S

Mt. Humboldt Gossia alaternoides (Brongn. & Gris) N. Snow (1618 m) Gossia angustifolia N. Snow, sp. nov. 22° 22°

100 km Nouméa

164°E 165° 166° 167°

Fig. 3. — Distribution maps of Gossia alaternoides (Brongn. & Gris) N. Snow and G. angustifolia N. Snow, sp. nov. Note: in all distribution maps, gray areas indicate ultramafic substrates,slash-hatching indicates dense humid forests, and dotted areas indicate protected areas; icons with “T” indicate type localities.

Petals stamens and style Gossia aphthosa (Vieill. ex Brongn. & Gris) N. Snow Unknown. (Figs 4D-G; 5; 13E-I)

Berry Austrobaileya 8: 180 (2010). — Eugenia aphthosa Brongn. & 6-8 × c. 7 mm, globular, base rounded, whitish to reddish; seeds 2. Gris, Bulletin de la Société botanique de France 13: 469 (1865). — Austromyrtus aphthosa (Brongn. & Gris) Burret, Notizblatt des Botanischen Gartens und Museums zu Berlin-Dahlem 15: 504 (1941). — Remarks Typus: New Caledonia. Grande Terre, South Prov., Collines de Gossia angustifolia sp. nov. is recognized by its narrowly elliptic Wagap, Vieillard 2172 (holo-, P[P00602542]; iso-, A[A00255453, leaves with slightly undulate (in sicco) margins and (usually) A00255454], B, BISH, G[G003410963, G00340964], L[3 sheets], dense but small oil glands on the lower surface. It can resemble MEL, P[P00602543], Z[Z-000050852]). G. ramiflora sp. nov. based on the slightly undulate leaf mar- Etymology. — Likely from the Greek aphtha, which is a reference gins and their overall shape, but the margins of G. ramiflora to thrush (= mouth ulcers), coupled with the Latin suffix osa (“full sp. nov. typically are significantly more undulate. It also can of”). Aphthosa thus presumably refers to the ring of corky tissue at resemble G. vieillardii, but that species has fewer oil glands on the base of the petioles. the lower leaf surface. Preliminary results suggest this species Distribution, habitat and phenology. — In three distinct regions is atypical in the genus by virtue of its basal placentation in a areas on Grande Terre (Fig. 5); in gallery forests and other humid to wet single locule (Fig. 4C). The laminar oil glands are less dense forests over ultramafics, schists, micaschists, and gneiss, 250-500 m. on one specimen (Vandrot 597). Flowering August through February; fruiting September through June.

ADANSONIA, sér. 3 • 2020 • 42 (7) 139 Snow N.

Description Calyx Trees or shrubs Lobes 4 or 5, 1-2 mm, broadly triangular, sparsely sericeous 2-8 m. above, glabrous to sparsely sericeous below, green.

Branchlets Petals Terete to compressed; internodes > 5 mm. c. 5 × c. 4 mm, glabrous above and below, ciliate on margins.

Leaves Filaments Coriaceous, 2 per node; petioles 2-4.8 mm, flat to terete; blades 5-8 mm; anther sacs 0.4-0.5 mm. (8.5-)12.0-24.0 × (3.5-)6.6-13.0 cm, elliptic to broadly elliptic, base cordate (and sometimes amplexicaulous) to broadly Style rounded, surface flat, margin flat to somewhat revolute at 6-7 mm, glabrous to sparsely sericeous at base. edges, apex obtuse to somewhat acute, midnerve above sulcate, secondary veins flush above, oil glands of lower surface Berry dense but indistinct. 8-9 × 8-9 mm, globular, base rounded, green maturing becoming dark purplish to blackish; seed number unknown. Inflorescence 1.5-6 cm, of monads, triads, or few-flowered racemes, axillary Remarks or clustered on naked branches; pedicels 0.3-1.5 cm; extrafloral Gossia aphthosa can be distinguished among New Caledonian bracts lacking or scale-like. congeners by its relatively large, flat, and broadly rounded to cordate leaf bases and frequently cauliflorous inflorescences. The Bracteoles nominal subspecies has a prominent ring of corky tissue at the (1.5-)3-4 mm, very narrowly ovate or elliptic to very narrowly base of the petioles, which is unique in the genus (e.g., Snow obovate, sparsely sericeous. et al. 2003). The three subspecies are based on differences in eco-geography and non-fixed, but mostly consistent differences Hypanthium in morphology. The inflorescence branches are said to be red- c. 1.5 × 1.5 mm, campanulate, surface smooth, glabrous to dish on some specimens (e.g., MacKee 13229) and the leaves densely sericeous; ovary apex glabrous. are said to emerge pinkish (MacKee 4427).

Key to the subspecies of Gossia aphthosa (Vieill. ex Brongn. & Gris) N. Snow 1. Base of petiole surrounded by prominent ring of corky tissue; hypanthium often silvery-sericeous (northern late Cretaceous micaschistes and gneiss, Massif du Panié, Northern Province) ...... Gossia aphthosa subsp. aphthosa — Base of petiole without corky swellings; hypanthium mostly glabrous (northern Central Range basement Meso- zoics ands schistes) ...... 2 2. Inflorescences 5-8 cm; leaf blades stiffly coriaceous, base strongly cordate and clasping; north-central humid forests ...... Gossia aphthosa subsp. longipedunculata N. Snow & Munzinger, subsp. nov. — Inflorescence less than 6 cm; leaf blades coriaceous, base rounded or cordate and clasping; southeast humid forests over ultramafics...... Gossia aphthosa subsp. austro-orientalis N. Snow & K. Gandhi, subsp. nov.

Gossia aphthosa subsp. aphthosa Remarks (Figs 4D-G; 5) A reliable diagnostic character is the corky swellings surround- ing the base of the petioles (Fig. 4D-E, G). The hypanthium of some (e.g., MacKee 28444) can be prominently sericeous; the Specimens examined. — New Caledonia. North Prov. Pla- teau du Col des Roussettes route forestière près de Riv Ouen leaf blades of some appear to be only thinly coriaceous (e.g., – Sieu, 21°25’45.19”S, 165°27’15.01”E, 29.IX.1964, Blanchon Munzinger 3084). 994 (NOU[NOU029700], P[P00462860]); Touho, Ponandou, 20°49’12.00”S, 165°13’45.12”E, 15.V.1973, MacKee 26722 (NOU[NOU053234]); Tiwaka: Pente Sud, Inédète, 250 m, 30.III.1974, MacKee 28444 (NOU[NOU029696], P[P06668973]); Gossia aphthosa subsp. austro-orientalis Ponandou, le Captage, 27.X.2005, Munzinger 3084 et al. N. Snow & K. Gandhi, subsp. nov. (P[P05121899], WELTU). (Fig. 5)

Representative herbarium specimen. — P05121899. Differing from Gossia aphthosa subsp. longipedunculata N. Snow & Munzinger, subsp. nov. by its generally shorter inflorescences, leaf Distribution, habitat and phenology. — Central and northeast blades of somewhat thinner texture, and bases of leaves that are not parts of Grande Terre; over micaschistes and gneiss, 250 m. as tightly clasping the stem.

140 ADANSONIA, sér. 3 • 2020 • 42 (7) Revision of New Caledonian Gossia N. Snow & Guymer

A B

I J

D H

Fig. 4. — A-C, Gossia angustifolia N. Snow, sp. nov.: A, branch; B, inflorescence in part;C , flower with petals and stamens removed, and in longitudinal section at left; D-G, Gossia aphthosa (Vieill. ex Brongn. & Gris) N. Snow subsp. aphthosa: D, branch showing leaves with corky swellings at base of leaf and ramiflorous inflorescences; E, detail of corky swellings; F, detail of flower with reflexed petals; G, remnants of inflorescence in axil (with corky swellings) of dehisced leaf; H-J, Gossia bourailensis N. Snow, sp. nov.: H, branch with inflorescences; I, detail of flower bud; J, detail of flower with petals removed showing calyx lobes, staminal ring, hairy ovary apex, and style with proximal pubescence. Vouchers: A-C, MacKee 33461 (NOU); D, F, MacKee 28444 (P); E-G, MacKee 26722 (NOU); H-J, Balansa 1516a (P); K, Veillon 5958 (NOU); J, MacKee 5328 (P). Scale bars: A, D, E, G, H, 2 cm; B, 5 mm; C, F, I, J, 3 mm.

ADANSONIA, sér. 3 • 2020 • 42 (7) 141 Snow N.

Typus. — New Caledonia. Grande Terre, South Prov., Rivière ee 25749 (NOU[NOU053235]); ibid. loco, 21°11’00.99”S, Bleue, 25.X.1974, M. Schmid 5180 (holo-, P[P00462872]; iso-, 165°17’48.98”E, 28.IV.1973, MacKee 26628 (NOU[NOU053235, [NOU[NOU029695], P[P00462871]). NOU053038); Haute Tchamba, 21°0’55”S, 165°15’6”E, 10.XI.2002, Munzinger 1456 et al. (MO[MO-04767308], NOU[NOU002675], Paratypi. —New Caledonia. Grande Terre, South Prov., Forêt P[P00354535], WELTU). du Mois de Mai, 22°07’06.99”S, 166°38’48.01”E, 19.XII.2006, Barrabé et al. 431 (NOU[NOU016015], MO[MO-6012551], Representative herbarium specimen. — P00354535. P[P04776185]); Forêt du Mois de Mai, 13.VI.1951, Baumann- Bodenheim 14066 (P[P00462856]); ibid. loco, 24.VI.1951, Baumann- Etymology. — In reference to the relatively long peduncles oc- Bodenheim 14182 (A, P[P00462857], Z); ibid. loco, 24.VI.1951, curring on many specimens. Baumann-Bodenheim 14206 (A, P[P00462859], Z); Vallée de Thy, Distribution, habitat and phenology 22°07’06.99”S, 166°32’12.84”E, 28.VI.2003, Biffin 112 & Craven . — Central Grande (CANB[CANB799723, n.v.], P[P02090975]); Thy Upper Western Terre, near Ponérihouen, Haute Tchamba, and Haute vallée de Road forest, 340 m, 5.II.1980, Brinon 692 (P[P00462861]); St. Louis, l’Amoa (Fig. 5) in humid forests on volcanic-sedimentary soils, 600 m, 14.XI.1949, MacDaniels 2216 (P[P00462864]); Mt. Koghi, 300-900 m. Flowering confirmed August through April; fruiting slope towards Vallée de Thy (St. Louis), 400-500 m, 21.IV.1956, October and November. MacKee 4427 (A, L, P[P00462866]); ibid. loco, 21.IV.1956, MacKee 4446 (P[P00462867]); Forêt Desmazures, 13.IV.2005, Munzinger Remarks 2767 (NOU[NOU07908], P[P04827941, P04827945]); Haut This subspecies is recognized most easily by its large (8-12 mm) Yaté, Rivière Bleue, 300 m, Schmid 5269 (NOU[NOU029684, NOU029688], P[P00462873]); Haute Yaté: Forêt du Mois de petals. (An unpublished manuscript name of mine for some Mai, 200 m, 25.XI.1982, Suprin 2259 (P[P00462874]); ibid. loco, specimens was Gossia “grandiflora”.) The inflorescences often 8.III.1966, Veillon 660 (NOU[NOU029689], P[P00462875]). — are densely fasciculate in leaf axils or on branches and stems. Ab loco. 1868-1872, Balansa 88 (A, P[P00462854, P00462855]). It can also resemble G. kuakuensis, but that species has larger Representative herbarium specimen. — P04827941. flowers and a slightly ribbed to rugose hypanthium.

Etymology. — From the Latin australis (south) and oriens (east), in reference to its more southerly and easterly distribution on Grande Gossia bourailensis N. Snow, sp. nov. Terre relative to the other subspecies. (Figs 4; 6) Distribution, Habitat and Phenology. — Southern part of Grande Terre; in dense humid forests or riparian forests over ultra- Resembling Gossia diversifolia but differing by its terete branchlets mafics,c. 150-660 m. Flowering March through June; fruiting April and the leaf blade, which is conduplicate only at its base. through October. Typus. — New Caledonia. Grande Terre, South Prov. Bourail, dans les bois, III.1869, Balansa 1516a (holo-, P[P00724290]; iso-, Remarks BISH[fragment], P[P00402718, P00402755]). The subspecies varies from being shrubs from 3 m to trees Paratypi. — New Caledonia, Grande Terre, South Prov. Bourail, 1868- to 10 m. The fruit is said to transition from green to white, 1970, Balansa 416a (Z[Z-000050855]); Peya (Bourail), 14.IV.2011, followed by reddish to dark purple. The pulp of the mature J.-P. Butin 58 (P[P02089712]); Montagne des Sources, 21°08’09.96”S fruit is said to be sour (MacKee 4446). 166°36’10.80”E, 19.II.1982, Veillon 5958 (NOU[NOU0534329]). Representative herbarium specimen. — P00724290.

Etymology. — Derived from the town of Bourail. Gossia aphthosa subsp. longipedunculata N. Snow & Munzinger, subsp. nov. Distribution, habitat and phenology. — A poorly understood (Figs 5; 13E-I) species as presently known and collected only from the vicinity of Bourail (in 1869) and more recently (1982) from Montagne des Sources (Fig. 6); in woods, c. 50 m. Flowering February and March; Differing fromGossia aphthosa subsp. austro-orientalis N. Snow & K. fruiting through August. Gandhi, subsp. nov. by its somewhat longer inflorescences (5-8 cm vs less than 6 cm in G. a. subsp. austro-orientalis N. Snow & K. Description Gandhi, subsp. nov.), more stiffly coriaceous leaf blades, and different eco-geography. Also resembling Gossia colnettiana, but that species Shrubs has rounded to cuneate leaf bases (vs cordate-clasping leaf bases in 4-5 m. G. a. subsp. austro-orientalis N. Snow & K. Gandhi, subsp. nov.). Branchlets Typus. — New Caledonia. Grande Terre, North Prov., Haute Terete; internodes > 5 mm. Amoa, 21°00’02.98”S, 165°14’31.99”E, 300-400 m, 7.IX.1968, MacKee 19502 (holo-, P[P05094105]; iso-, NOU[NOU029685]). Leaves Paratypi. — New Caledonia. Nékoro, prop. P. Johnson, 1.IV.2004, Coriaceous, 2 per node; petioles 1-3 mm, rounded; blades Dagostini 654 (NOU[NOU001193); Haute Vallée de l’Amoa, 2.0-3.7 × 1.3-2.3 cm, ovate, base rounded to cordate, surface 21°00’02.98”S, 165°14’31.99”E, 12.VIII.1965, MacKee 13229 flat to slightly wavy but somewhat conduplicate above petiole, (BISH, MO[MO-04777159], NOU[NOU023797], P[P00462868, P00462869, P00462870]; ibid. loco, 300 m, 28.IV.1968, MacK- margin flat to slightly undulate, apex broadly acute to obtuse, ee 18698 (NOU[NOU029687], P[P02089984]); Ponérihouen, midnerve above flush, secondary and tertiary veins slightly raised 21°11’00.99”S, 165°17’48.98”E, 700-900 m, 6.XI.1972, MacK- throughout, oil glands of lower surface sparse and indistinct.

142 ADANSONIA, sér. 3 • 2020 • 42 (7) Revision of New Caledonian Gossia N. Snow & Guymer

165°E 166° N

Mt. Panié Ouvéa (1629 m) Island

21°S Koné 21°S

Gossia aphthosa (Brongn. & Gris) N. Snow subsp. aphthosa Mt. Humboldt Gossia aphthosa (Brongn. & Gris) N. Snow (1618 m) subsp. austro-orientalis N. Snow & K. Gandhi, subsp. nov.

22° Gossia aphthosa (Brongn. & Gris) N. Snow 22° subsp. longipedunculata N. Snow & Munzinger, subsp. nov.

100 km Nouméa

164°E 165° 166° 167°

Fig. 5. — Distribution map for subspecies of Gossia aphthosa (Brongn. & Gris) N. Snow.

Inflorescence Style 1.4-1.8 cm, of monads, terminal, axillary, or from naked 4-4.5 mm, sparsely sericeous to tomentose throughout. internodes; extrafloral bracts absent or scale-like. Fruits Bracteoles Green 4-5.5 × 4-5 mm, globular, base rounded, mature color 0.5-1 mm, narrowly ovate to ovate, glabrous to sparsely sericeous. unconfirmed; seed number 1.

Hypanthium Remarks 1.5-2.2 × 2.5 mm, obconic to campanulate, surface smooth, Gossia bourailensis sp. nov. resembles G. diversifolia but has glabrous; ovary apex hairy. terete branchlets, a somewhat narrower petiole, and the base of the leaf blade is more or less conduplicate. It also resembles Calyx G. conduplicata sp. nov., which has much more pronounced Lobes 4 or 5, 0.5-2 mm, broadly rounded, glabrous above folding of the leaf blades, and Gossia katepahiensis sp. nov., and below, green. which has a more broadly rounded leaf apex and glabrous ovary. Some petioles of G. bourailensis sp. nov. are twisted Petals and bent away from the axis of the midrib, but this is a subtle 4 (or irregularly 5-8), 4-5 × 2.8-4.5 mm, glabrous above and character. Some flowers of Balansa 416a have eight petals, below, minutely ciliate on margins. which probably is a developmental anomaly. The distribution as currently understood, from the type specimen in 1869 Filaments near the west-central part of Grande Terre and Montagne 3.5-5 mm; anther sacs 0.3-0.5 mm. des Sources in 1982 is anomalous; further study is needed.

ADANSONIA, sér. 3 • 2020 • 42 (7) 143 Snow N.

165°E 166° N

Mt. Panié Ouvéa (1629 m) Island 21°S 21°S Koné

Gossia bourailensis N. Snow, sp. nov. Gossia colnettiana (Guillaumin) N. Snow Mt. Humboldt (1618 m) Gossia conduplicata N. Snow, sp. nov. Gossia diversifolia (Brongn. & Gris) N. Snow 22° 22°

100 km Nouméa

164°E 165° 166° 167°

Fig. 6. — Distribution maps for Gossia bourailensis N. Snow, sp. nov., G. colnettiana (Guillaumin) N. Snow, G. conduplicata N. Snow, sp. nov. and G. diversifolia (Brongn. & Gris) N. Snow.

Gossia clusioides (Brongn. & Gris) N. Snow Branchlets (Figs 7-11) Round to compressed; internodes > 5 mm.

Austrobaileya 8: 180 (2010). — Eugenia clusioides Brongn. & Gris, Leaves Bulletin de la Société botanique de France 12: 180 (1865). — Austromyrtus clusioides (Brongn. & Gris) Burret, Notizblatt des Coriaceous, 2 per node; petioles 3.5-20 mm, slightly sulcate; Botanischen Gartens und Museums zu Berlin-Dahlem 15: 503 blades 7.0-18.5 × 2.5-12.0 cm, narrowly elliptic to broadly (1941). — Typus: New Caledonia. Grande Terre, South Prov., elliptic or obovate, base cuneate or rounded, surface flat to Deplanche 525 (holo-, P[P00602545]). strongly bullate, margin flat to revolute, apex obtuse to some- Etymology. — Derived from the generic name Clusia L. (Clu- times retuse, midnerve above strongly sulcate, secondary veins siaceae) given the resemblance of its leaf blades to many species flush to impressed above, oil glands of lower surface sparse to of that genus. common but indistinct.

Distribution, habitat and phenology. — Across much of Grande Terre (Fig. 7); in humid or gallery forests over micaschistes, Inflorescence graywackes, or ultramafics, 10-900 m. Flowering December to June; 90-160 mm; of monads, cymes, racemes, or panicles, terminal fruiting January to November. or axillary, pedicels 0.7-2.0 cm; extrafloral bracts leafy and pronounced when present but soon deciduous. Description Trees or shrubs Bracteoles (0.5-)3-9(-15) m tall. 6-7 mm, narrowly ovate to ovate, sparsely sericeous.

144 ADANSONIA, sér. 3 • 2020 • 42 (7) Revision of New Caledonian Gossia N. Snow & Guymer

165°E 166° N 100 km

Mt. Panié Ouvéa (1629 m) Island 21°S 21°S Koné

Gossia clusioides (Brongn. & Gris) N. Snow subsp. ploumensis (Däniker) N. Snow, comb. et stat. nov. Gossia clusioides subsp. callmanderiana N. Snow, subsp. nov. Gossia clusioides subsp. clusioides Mt. Humboldt Gossia clusioides subsp. rembaiensis N. Snow, subsp. nov. (1618 m) Gossia clusioides subsp. taomensis N. Snow, subsp. nov.

22° Gossia clusioides subsp. avanguiensis N. Snow, subsp. nov. 22° Gossia clusioides subsp. bleuensis N. Snow, subsp. nov. Gossia clusioides subsp. maoyana N. Snow, subsp. nov. Gossia clusioides subsp. tiebaghiensis N. Snow, subsp. nov. Nouméa

164°E 165° 166° 167°

Fig. 7. — Distribution map for subspecies of Gossia clusioides (Brongn. & Gris) N. Snow.

Hypanthium Remarks 4-5 × 6-11 mm, campanulate, surface smooth, sparsely seri- Gossia clusioides has large flowers and small leafy bracts in the ceous to sericeous (trichomes sometimes reddish at base); inflorescences that typically are soon caducous; the species ovary apex glabrous. also can be diagnosed by the strongly sulcate midveins on the upper leaf surface. It is one of several species in which the Calyx epidermis begins to crack into dark, irregularly rectangular Lobes 5, 2.5-4.5 mm, broadly ovate to rounded, sericeous flakes during the second year of growth. above (sometimes densely so), sparsely sericeous below, green. Herbarium labels indicate that some members are slender and somewhat sparsely branched. Specimens with large (non- Petals bullate) leaves might be confused with of G. aphthosa, but 9-11.5 × 7-9 mm, glabrous above, sericeous-tomentose below the flowers and fruits of G. clusioides are considerably larger. (trichomes reddish at base). The species has considerably more variation in leaf morphology than others of Gossia in New Caledonia. As treated Filaments here, Gossia clusioides comprises nine subspecies that are 6-8 mm; anther sacs 0.6-1.0 mm. separated based primarily on differences in leaf dimensions and texture (smooth versus bullate), edaphic preferences, Style and often (but not always) geographic isolation. Some sites, Up to 9 mm, glabrous. however, have more than one subspecies. Much additional fieldwork is needed to further test the distinctness of the Fruits subspecies and to better understand their differences and 7-17 × 7-15 mm, globular, base rounded to tapered, dark distributions, especially given that these entities may be bluish-black; seeds 1-3. undergoing speciation.

ADANSONIA, sér. 3 • 2020 • 42 (7) 145 Snow N.

Key to the subspecies of Gossia clusioides (Brongn. & Gris) N. Snow 1. Leaf blades moderately to strongly bullate ...... 2 — Leaf blades flat (sometimes slightly bullate in subspp.clusioides and taomensis) ...... 5 2. Mature leaves (6.5-)7.0-12.5 cm wide ...... Gossia clusioides subsp. ploumensis (Däniker) N. Snow, comb. et stat. nov. — Mature leaves 3.5-6.5 cm wide ...... 3 3. Leaves ovate to elliptic or broadly elliptic ...... Gossia clusioides subsp. maoyana N. Snow, subsp. nov. — Leaves narrowly elliptic ...... 4 4. Leaves 4-7 times longer than wide ...... Gossia clusioides subsp. avanguiensis N. Snow, subsp. nov. — Leaves 2-3(-4) times longer than wide ...... Gossia clusioides subsp. bleuensis N. Snow, subsp. nov. 5. Leaves mostly elliptic to broadly elliptic or obovate (rarely narrowly elliptic in subsp. clusioides) ...... 6 — Leaves narrowly elliptic ...... 7 6. Leaf blades thickly coriaceous, upper surface often glossy, abaxially secondary veins often somewhat distinct, margin often strongly recurved nearly throughout; southwestern coast over ultramafics ...... Gossia clusioides subsp. callmanderiana N. Snow, subsp. nov. — Leaf blades coriaceous, upper surface matte, abaxial secondary veins flush or distinct, margins flat to slightly recurved; across much of Grande Terre ...... Gossia clusioides subsp. clusioides 7. Leaves obovate to elliptic, apex obtuse or (usually at least slightly) retuse ...... Gossia clusioides subsp. rembaiensis N. Snow, subsp. nov. — Leaves narrowly elliptic, apex obtuse to broadly acute ...... 8 8. Petioles 2-7 mm long; leaves 3-6 cm long ...... Gossia clusioides subsp. taomensis N. Snow, subsp. nov. — Petioles 8-20 mm long; leaves 9.5-17 cm long ...... Gossia clusioides subsp. tiebaghiensis N. Snow, subsp. nov.

Gossia clusioides subsp. avanguiensis N. Snow, subsp. nov. resent this subspecies; its developing leaves remain stramineous (Figs 7; 11A) to light maroon until they reach full size and become green.

Similar to Gossia clusioides subsp. bleuensis subsp. nov. but with leaf blades 4-7 × longer than wide (vs 2-3[-4] × longer in G. c. subsp. bleuensis subsp. nov.), more strongly bullate, and occurring near the west central Gossia clusioides subsp. bleuensis N. Snow, subsp. nov. coast at about 400 m (vs subsp. bleuensis subsp. nov. in the southeast). (Figs 7; 8A; 10B) Typus. — New Caledonia. Grande Terre, North Prov. Poya: Avan- Leaf blades strongly bullate, 4.0-8.0(-11.5) cm, narrowly elliptic. gui, 400 m, 3.VIII.1976, in low forest on rocky serpentine slope, The inflorescence can be up 8.5 cm long, exceeding adjacent leaves. MacKee 30824 (holo-, P[P02089852]). The growth form in profile often is slender. Paratypus. — New Caledonia. Grande Terre, North Prov. Poya: Avan- Typus. — New Caledonia. Grande Terre, South Prov., Rivière Bleue gui, 2.I.1976, MacKee 30629 (NOU[NOU029543], P[P02089853]). reserve, forested slopes 300 m, 30.VII.1981, McPherson 4001 (holo-, P[P00402735]; iso-, BISH[fragment], MO, NOU[NOU030915], Representative herbarium specimen. — P02089852. PTBG, WELTU). Etyomology. — After Avangui, the type locality. Paratypi. — New Caledonia. Grande Terre, South Prov., Rivière Bleue, 22.II.1951, Baumann-Bodenheim 10873 (P[P00402729]); Bon Secours, Distribution, ecology and phenology. — Poya (Avangui) (Fig. 7); Baumann-Bodenheim 11847 (P[P00402730]); Rivière Bleue, Mois de in low forests over ultramafics, to 400 m. Flowering confirmed only Mai, 23.VI.1951, Baumann-Bodenheim 14045 (MO[MO-5163034], for March; fruiting confirmed only for January. P[P00402727]); Forêt Faux Bon Secours, 9.V.1981, MacKee 39042 (NOU[NOU029540], P[P02089862, P02089863]). Remarks MacKee sometimes returned to the same specimen to collect Representative herbarium specimen. — P02089863. flowers and fruits at different times, and the two specimens Etymology. — From its occurrence in the vicinity of Rivière Bleue. cited likely are from the same tree (P. Lowry pers. comm. 2015). Distribution, habitat and phenology. — Southeast Grande The unvouchered photo (Fig. 11A), which almost assuredly Terre (Fig. 7); forested slopes over ultramafics, 300-450 m. Flower- is this taxon, is said to be from Näräja, between Ponérihouen ing May and June; fruiting confirmed only for July to December. and Ouaté, but I have been unable to confirm that location. This subspecies, as presently known from vouchered collec- Remarks tions, is a dense, multicaulous shrub of c. 1 meter. However, Further fieldwork may reveal this as being the same taxon asG. the non-vouchered digital images (Fig. 11A; see also http:// a. subsp. avanguiensis subsp. nov., which is diagnosed primar- endemia.nc/flore/fiche6365) of a larger shrub appears to rep- ily by its longer leaves, but which occurs much farther north.

146 ADANSONIA, sér. 3 • 2020 • 42 (7) Revision of New Caledonian Gossia N. Snow & Guymer

A B C

F H

Fig. 8. — Typical leaf morphology for most subspecies of Gossia clusioides (Brongn. & Gris) N. Snow: A, G. c. subsp. bleuensis N. Snow., subsp. nov., with narrowly elliptic, bullate blades; B, G. c. subsp. maoyana N. Snow, subsp. nov., blades relatively narrow and bullate but longer than those of subsp. bleuensis; C, D, G. c. subsp. callmanderiana N. Snow, subsp. nov., blades elliptic and flat; E, G. c. subsp. rembaiensis N. Snow, subsp. nov.; F, G. c. subsp. clusioides; G, G. c. subsp. subsp. taomensis N. Snow, subsp. nov.; H, G. c. subsp. tiebaghiensis N. Snow, subsp. nov. Vouchers: A, McPherson 4001 (BISH); B, Dawson 16603 (WELTU)); C, Jaffré 1275 (BISH); D, Virot 1537 (BISH); E, McPherson 5083 (BISH); F, MacKee 25608 (NOU); G, Veillon 4828 (NOU); H, McPherson 3323 (NOU). Scale bar: 2 cm.

Gossia clusioides subsp. callmanderiana N. Snow, subsp. nov. Paratypi. — New Caledonia. Grande Terre, South Prov., Route (Figs 7; 8C, D; 10A) de Prony, 17.XII.1973, Jaffré 1197 (NOU[NOU029545], P[P00402724]); Région du Col de Plum, II.1974, Jaffré 1275 (BISH[fragment], NOU[NOU029546], P[P00402731]); Plum, Leaf blades flat, thickly coriaceous, and often with a glossy adaxial 18.V.1985, MacKee 42596 (NOU[NOU029581], P[P05094060]); surface; margin becoming recurved more or less throughout; secondary Plum, 30 m, 31.III.1986, MacKee 43056 (P[P02089854]); Au-dessus abaxial veins often prominent and projecting slightly. de la Baie N’go, 4.II.1982, Veillon 4813 (NOU[NOU029547], P[P05094308]); Rive Nord de la Baie N’go, 28.IV.1945, Virot Typus. — New Caledonia. Grande Terre, South Prov., Baie 1537 (BISH, P[P00402721, P00402748, P00402749], WELTU). Ngo, 28.VII.1983, Suprin 2221 (holo-, P[P00402738]; iso-, NOU[NOU053238], P[P00402752]). Representative herbarium specimen. — P02089854.

ADANSONIA, sér. 3 • 2020 • 42 (7) 147 Snow N.

Etyomology. — In recognition of Martin Wilhem Callmander Gossia clusioides subsp. maoyana N. Snow, subsp. nov. (1975-) of the Conservatoire et Jardin botaniques de la Ville de (Figs 7; 8B) Genève (G), for his many collections and editorial commitment to expedite the publication of taxonomic novelties from New Trees or shrubs 3-4 m. Leaf blades slightly bullate, ovate to elliptic Caledonia (e.g., Snow et al. 2016a, b). or broadly elliptic.

Distribution, ecology and phenology. — Vicinity of Baie Typus. — New Caledonia. Grande Terre, North Prov., Nord Massif Ngo and Plum (Fig. 7); in maquis over ultramafics, up to 50 m. Ignambi, 600 m, 28.XI.1967, Veillon 1514 (holo-, P[P00758089]; Flowering March through July; fruiting confirmed only for February. iso-, NOU[NOU030918]).

Paratypi. — New Caledonia. Grande Terre, North Prov., Mé Maoya, Remarks 780 m, 20.XI.1992, Jaffré 3196 (NOU[NOU029543]); Mé Maoya, Athough somewhat subtle on herbarium specimens, the 500 m, 30.XI.1992, Dawson WELTU 16603 (BISH[fragment], leaf texture is thicker and the sheen of the adaxial surface WELTU). — Ab loco., Balansa 1490 (P[P00402745]). is more glossy (reflective) than members of G. a. subsp. Representative herbarium specimen. — P00758089. bleuensis subsp. nov. and G. a. subsp. clusioides, which also occur in the southeast. Etymology. — In reference to collections from Mé Maoya. Distribution, habitat and phenology. — Mé Mayoa and Ig- nambi (Fig. 7); in dense but often low stature forests over ultrama- Gossia clusioides subsp. clusioides fics, 400-780 m. Flowering confirmed February to April; fruiting (Figs 7; 8F) January through November.

Specimens examined. — New Caledonia. Grande Terre. Remarks North Prov. Col de Poro (Bota Mere), 10.II.2004, Dagostini & Rigault 792 (NOU[NOU002967], P[P05121901]); Ponéri- Additional sampling may reveal this to be the same as the houen: Pente est du Mont Aoupinié, 500-700 m, 27.VI.1972 , nominal subspecies, although the latter occurs in the southeast. MacKee 25608 (BISH, NOU[NOU029590], P[P00402732]); Some specimens, including the holotype, were annotated pre- Haute Néaoua-Ouen-Sieu, 500-600 m, 12.II.1970, MacKee viously with a manuscript infraspecific epithet of “orientalis”. 21835 (NOU[NOU029589]); Kopeto, crête est du Mont Vert, 700 m, 24.VI.1973, MacKee 26893 (P[P02089844]); Koné, Massif Koniambo, Creek Confiance, rive gauche, 21°02’07.85”S, 164°48’42.33”E, 100 m, 21.IX.2016, Muzinger 7875 (legit. Gossia clusioides (Brongn. & Gris) N. Snow Scoptera) (MPU[MPU640696]). — South Prov., Bassin du subsp. ploumensis (Däniker) N. Snow, comb. et stat. nov. Boulari, 17.V.1869, Balansa 1490a (A, P[P00402723], Z); Col (Figs 7; 9; 10D) de Mouirange, forêt Desmazure, 8.III.2005, Dagostini 1040 (NOU[NOU009286]); Ngo Col, 100 m, 22.X.2004, Dagostini Differing from other subspecies by the relatively large, elliptic, and 1296 (NOU[NOU019100]); À Yaté, 20.III.1916, Franc 2084 moderately to strongly bullate leaf blades. (A, BISH, NOU[NOU029555], P[P00402728]); Kuébini, 2 m, 21.VI.1968, MacKee 19007 ([NOU029583], P[P02089849]); Austrobaileya 8: 180 (2010). — Eugenia ploumensis Däniker, Kuébini, 5.III.1971, MacKee 23393 (NOU[NOU029582], Beiblatt zur Vierteljahrsschriftder Naturforschenden Gesellschaft in P[P02089856, P02089857]); Yaté, Touaourou, 5 m, 21.XII.1986, Zürich, Beibl. 19, 78: 298 (1933). — Austromyrtus ploumensis MacKee 43385 (NOU[NOU029579], P[P00402720]); Mt Naka- (Däniker) Burret, Notizblatt des Botanischen Gartens und Museums zu da, c. 900 m, VI.1979, Morat 6411 (P[P00462887]); Plaine de Berlin-Dahlem 15: 504 (1941). — Typus: New Caledonia. Grande Kanala, 1861-1867, Vieillard 2597 (P[P00402750]); Bois de la Terre, South Prov., Ploume, 9.II.1926, A.U. Däniker 2807 (holo-, Plaine, Kanala, Vieillard 2612 (P[P00402751]); Neuméníe, Mé Z[Z-000016108]; iso-, Z[Z-000016109]). Ouébo, c. 350 m, 18.XII.1981, Veillon 4772 (NOU[NOU029554], P[P00402739], WELTU); Riv. Tara, 2.5-3 km N of Touaourou, Distribution, habitat and phenology. — Across much of 10-20 m, 18.VIII.1968, Webster & Hildreth 14881 (DAV, GH, Grande Terre (Fig. 7); in forests; c. 300-600 m. Flowering August NSW, P). — Ab loco. Balansa 1490 (P[P05093917]); Petit 166 through March; fruiting confirmed from December through June. (P[P00402737]). Specimens examined. — New Caledonia. Grande Terre, North Representative herbarium specimen. — P02089856. Prov. Silva Peala, ad occidentem Pouebo, 500-600 m, 6.V.1968, Bernardi 12606 (P[P00402719]; Mount Mandjelia and slopes, Distribution, habitat and phenology. — Across much of 540 m, 20°24’19”S, 164°31’01”E, Craven 15027 et al. (CANB, Grande Terre, sometimes near coast in southeast (Fig. 7); in n.v.; P[P02091556]); Forêt de Tende (Haute Diahot), exploita- maquis over ultramafics, 5-900 m.Flowering February to Sep- tion forestière Frouin, 500-600 m, 12.XI.1967, MacKee 17534 tember; fruiting February to December. (NOU[NOU029693], P[P02088630]); Forêt de Tende (Haute Diahot), 500-600 m, 17.IX.1967, MacKee 17550 (P[P02089850]); Haute Diahot: Exploitation Forestière, Frouin, Forêt Tendé, Remarks 31.III.1969, MacKee 20453 (NOU[NOU05323]); Haut Diahot, My concept of this subspecies has shifted significantly Tendé, 600 m, 30.VI.1982, MacKee 40567 (MO[MO-6751044], through time, rendering as outdated many prior annota- NOU[NOU029690], P[P02089872]); Road leading up to tions (some dating back to 2003), including some that Mt Mandjélia from Ouégoa, c. 650 m, 11.VIII.2003, Snow 9215 et al. (ASU, BISH, BRI, CANB, KSP[KSP016939], MO, were annotated as type specimens. Specimens at the NOU[NOU00683], NY, P[P00459485], WELTU). — South southern tip of Grande Terre can have narrowly elliptic Prov. Mt Mi (= Do Mt sentier Ouanémie [Hopkins & Bradford leaves (MacKee 23393). 2009: appendix 2]), 20.II.1869, Balansa 1490a (P[P00402725,

148 ADANSONIA, sér. 3 • 2020 • 42 (7) Revision of New Caledonian Gossia N. Snow & Guymer

Fig. 9. — Gossia clusioides (Brongn. & Gris) N. Snow subsp. ploumensis (Däniker) N. Snow, comb. et stat. nov., showing peeling and cracking bark of branchlets. Voucher: Snow 9215, BISH (sheet 3 of 3). Scale bars: A, C, 2 cm; B, 1 cm.

P06669014]); Entre St Louis et Ounia, 15.I.1869, Balansa Remarks 1490bis (P[P00402726, P00402753, P00402754]; Piste du pont This subspecies has large, elliptic and moderately to strongly des Japonais, 166°42’40”E, 22°15’31”S, 4.II.2007, Munzinger 4119 et al. (MO n.v., NOU n.v., P[P05321122, P05321125, bullate leaf blades. It occurs from the north to south near P05321126], WELTU). the coasts, but I am unable to discern consistent differences between the northern and southern populations. Previously Representative herbarium specimen. — P05321125. annotated specimens of this subspecies may bear the manu-

ADANSONIA, sér. 3 • 2020 • 42 (7) 149 Snow N.

Fig. 10. — Living specimens of Gossia clusioides: A, G. c. subsp. callmanderiana N. Snow, subsp. nov. vel aff.; B, G. c. subsp. bleuensis N. Snow, subsp. nov. vel aff.; C, G. c. subsp. ploumensis (Däniker) N. Snow, comb. et stat. nov.; D, G. c. subsp. ploumensis comb. et stat. nov. vel aff., but with larger and more prominently bullate leaves than normal and possibly representing an undescribed taxon. Vouchers: A, ©Hervé Vandrot, taken at Cascade la Pandanas-Koniambo in 2010 (http://endemia.nc); B, © Christian Létocart, taken at Yaté; C, ©Daniel & Irène Létocart, taken at Napoérédjeine in 2007; D, ©Jean-Louis Ruiz, taken at Pont des Japonais in 2005 (http://endemia.nc). script names Gossia “grandibullata” or Gossia clusioides subsp./ (NOU[NOU029580], P[P02089858]; Mt Nakada, vers 900 m, var. “grandibullata”. Unvouchered photos (Fig. 10D) of a VI.1979, Morat 6411 (P[P00462887]). possible specimen of this subspecies taken by J.-L. Ruiz near Representative herbarium specimen. — P00402736. the River de Pirogues showed honeybees (Apis sp.) pollinat- ing the flowers. Distribution, habitat and phenology. — Inland mountains on Grande Terre (Fig. 7); humid forested slopes over schistes or graywackes, 450-900 m. Flowering November through July; fruits not seen. Gossia clusioides subsp. rembaiensis N. Snow, subsp. nov. (Figs 7; 8E) Remarks Indicated as being common in some places locally (MacKee Leaf blades flat; differing from other subspecies by its attenuate base 44916). The flowers are said to have a spicy-fragrant smell of leaf and retuse apex. (McPherson 5083). MacKee (MacKee 29024) indicates the local name as being “tu”. Typus. — New Caledonia. Grande Terre, South Prov. Forested slopes near Mt Rembai, south of Col d’Amieu, c. 500 m, 7.XI.1982, McPherson 5083 (holo-, P[P00402736]; iso-, BISH[fragment], MO[MO-3218923, MO-2907911], NOU[NOU029586, Gossia clusioides subsp. taomensis N. Snow, subsp. nov. NOU67076]). (Figs 7; 8G; 11B)

Paratypi. — New Caledonia. Grande Terre, North Prov. Mt. Leaf blades flat to slightly bullate, narrowly elliptic to elliptic, Rembai, 450 m, 7.XI.1982, Dawson WELTU16591 (WELTU); 3-6 cm, less than 3 times as long as wide; petioles 2-7 mm long. Contrefort Nord-ouest du Mt. Canala, 700-800 m, 28.VII.1971, Inflorescences 3-7 cm long. MacKee 24011 (P[P02089847]); Ponerihouen, Pente est du Mont Aoupinie, 500 m, 27.VII.1974, MacKee 29024 (NOU[NOU029577], Typus. — New Caledonia. Grande Terre, North Prov., Mont P[P02089848]); Mont Pembai, 700 m, 31.V.1990, MacKee 44916 Taom: Crête Est, 1000 m, 13.VII.1979, MacKee 37148 (holo-,

150 ADANSONIA, sér. 3 • 2020 • 42 (7) Revision of New Caledonian Gossia N. Snow & Guymer

Fig. 11. — Living specimens of Gossia clusioides: A, G. c. subsp. avanguiensis N. Snow, subsp. nov. vel aff.; B, G. c. subsp. taomensis N. Snow, subsp. nov. Vouchers: A, ©Hervé Vandrot, taken at Naraja entre Ouaté et Ponérihoen (http://endemia.nc); B, Munzinger 3015; photo by J. Munzinger.

P[P00402734]; iso-, BISH, NOU[NOU029551, NOU029585], Etymology. — From the type locality. WELTU). Distribution, habitat and phenology. — Massif de Tiébaghi Paratypi. — New Caledonia. Grande Terre, North Prov. Base (Fig. 7); in creekside forests and degraded maquis over ultramafics, sud du Koniambo, Riv. Rouge, maquis sur terrain rocheux ser- c. 300-500 m. Flowering April; fruiting April through November. pentineux, 50 m, 23.III.1969, MacKee 20343 (P[P02089846]); Base sud de Koniambo (as « Koriambo »), Rivière Rouge, 50- 100 m, 15.IV.1972, MacKee 25301 (NOU[NOU029552], P[P02089983]; Taom, Mt. Homédéboa, 800-900 m, 16.X.1969, Gossia colnettiana (Guillaumin) N. Snow MacKee 20967 (NOU[NOU029587], P[P02089845]); Mont Pae- (Figs 6; 12A-D) oua, 900-1000 m, VI.1973, MacKee 26887 (NOU[NOU029584], P[P02089843]); Base du Koniambo, Rivière Pandanus, 3.X.2005, Austrobaileya 8: 180 (2010). — Eugenia colnettiana Guillaumin, Munzinger & Swenson 3015 (P[P04885463]); Massif du Taom, 900 m, Mémoires du Muséum national d’Histoire naturelle, Paris, sér. B, Bot. 24.III.1982, Veillon 4828 (NOU[NOU029585, NOU029551], 8: 242 (1962). — Typus: New Caledonia. Grande Terre, North WELTU). Prov., Mt Colnett, 1370 m, 13.IX.1951, Hürlimann 1973 (holo-, P[P00602541]; iso: Z[Z-000050851]). Representative herbarium specimen. — P00402734. Etymology. — Based on the type locality. Distribution, habitat and phenology. — Peaks of northwest coast (Koniambo and Massif du Taom; Fig. 7); in maquis over Distribution, habitat and phenology. — Mt Colnett and Mt ultramafics, c. 50-1000 m. Flowering confirmed only for March; Panié (Fig. 6); forests and forested slopes, presumably over meta- fruiting July through October. morphics, 1000-1500 m. Flowering September through November; fruiting confirmed for early May, but likely extending into mid June or longer. Remarks Evidently rare near the base of Koniambo. This subspecies Specimens examined. — New Caledonia. Grande Terre, North has leaves similar to, but of significantly smaller dimensions, Prov. Mt. Colnett, 1000 m, 27.X.2003, McPherson 18973 et al. (MO); Mt. Colnett, forested eastern slopes, 1000 m, 29.X.2003, than those of G. c. subsp. tiebaghiensis subsp. nov. McPherson 19021 (MO); ibid. loco, 29.X.2003, MacKee 19062 (MO); ibid. loco, 1250-1500 m, 31.X.2003, McPherson 19108 (BISH, KSP[KSP004707], MO, NOU[NOU004355]); Mont Panié, Gossia clusioides subsp. tiebaghiensis N. Snow, subsp. nov. 20°35’09.96”S, 164°46’14.99”E, 1300 m, 5.V.2007, Munzinger (Figs 7; 8H) 4337 et al. (NOU[NOU21487], P[P06668691]). Representative herbarium specimen. — P06668691. Leaf blades flat, 9.5-18.0 × 3.0-6.5 cm, narrowly elliptic. Inflorescence 5.0-6.5 cm. Description Typus. — New Caledonia. Grande Terre, North Prov., Dôme de Shrubs or trees Tiébaghi (Pente ouest), 400 m, 25.IV.1976, MacKee 31129 (holo-, 1-5 m. P[P00402733]; iso-, NOU[NOU030917]).

Paratypi. — New Caledonia. Grande Terre, North Prov., Pente Branchlets sud-ouest du Dôme de Tiebaghi, 300-500 m, 9.V.1966, MacKee Terete, surface smooth; internodes > 5 mm. 14935 (BISH, CHR n.v., MO, P[P02089851], WELTU n.v.); Massif de Tiébaghi, c. 12 air-km NW of Koumac, 8.XI.1980, McPherson Leaves 3323 (BISH, BRI n.v., MO[MO-3225918], NOU[NOU029550]). Coriaceous, 2 per node; petioles 2-2.8 mm, terete; blades 2.0- Representative herbarium specimen. — P00402733. 6.5 × 1.2-3.5 cm, elliptic, base rounded, surface flat, margin

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flat to somewhat revolute, apex obtuse to slightly acute, mid- Description nerve above sulcate, secondary veins above flush, oil glands Shrubs of lower surface common, sometimes faint. To 3 m.

Inflorescence Branchlets (2.0-)3.5-7.5 cm, of monads, simple cymes, or few-flowered Terete; internodes > 5 mm. racemes, terminal or axillary along upper nodes; pedicels 5-30 mm; extrafloral bracts leafy if present. Leaves Coriaceous, 2 per node; petioles 1.5-2.5 mm, flattened to broadly Bracteoles sulcate above; blades 2.0-3.0 × 1.2-2.0 cm, ovate, base rounded c. 3.5 mm, narrowly elliptic to narrowly obovate, glabrescent but conspicuously conduplicate above petiole (up to or more to sparsely sericeous. than half the length), surface moderately to strongly undulate, apex acute, midnerve above flush to slightly sulcate, secondary Hypanthium veins not raised above, oil glands absent from lower surface. Campanulate, 4-5 × 4.5-5 mm, sometimes slightly ribbed, more or less sericeous; ovary apex hairy. Inflorescence A monad or simple cyme, axillary and terminal, sometimes Calyx nodding apically; extrafloral bracts absent. Lobes 5, 3.5-5 mm, broadly ovate, sparsely ciliate on margins, glabrous above, glabrous to sparsely sericeous below, Bracteoles greenish-white (in vivo). c. 1 mm, narrowly elliptic, glabrous.

Petals Hypanthium 8-13 × 6-8 mm, glabrous above and below, minutely ciliate apically. c. 3 × 3 mm, campanulate, surface smooth, glabrous.

Filaments Calyx 8-14 mm; anther sacs 0.6-0.8 mm. Lobes 4(-5), 1.2-2.2 mm, broadly rounded, glabrous above and below, green. Style 11-16 mm, sparsely hairy at base. Petals Unknown. Fruit Not seen but said to be greenish when young (e.g., Munz- Filaments inger 4337). Not seen, estimate of c. 80.

Remarks Style The species closely resembles G. kuakuensis, but the latter has Not seen; ovary apex sparsely hairy. a glabrous hypanthium and occurs from 100-600 m in areas farther west or south, whereas G. colnettiana has a sericeous Fruit hypanthium and occurs at 1000 m or above. 4.5-5.5 × 4.5-5.5 mm, globular, base rounded, green (immature) to red (mature); seed 1.

Gossia conduplicata N. Snow, sp. nov. Remarks (Figs 6; 12E-G) This poorly known species is diagnosed by its strongly conduplicate leaves with undulate margins, and efforts to relocate Resembling Gossia diversifolia except that much to most of the blade it should be a high priority. The calyx lobes in fruit are re- itself is conduplicate, and it lacks the winged branchlets of G. diversifolia. flexed in the one known specimen, unlike the more common Typus. — New Caledonia. Grande Terre, North Prov., Poindimié, condition of ascending to erect lobes. The similar species 165°19’59.988’’E, 20°56’36.996’’S, 4.VIII.1967, MacKee 17222 G. diversifolia has winged branchlets. Gossia conduplicata sp. (holo-, P[missing]; iso-, P[P00781068], BISH[fragment], NOU[- NOU029074], WELTU). nov. differs from G. bourailensis sp. nov. by its more undulate leaf margin and narrowly acute leaf apex. The base of the Representative herbarium specimen. — P00781068. leaf blade in Gossia katepahiensis can be conduplicate, but Etymology. — In reference to the partially to strongly folded only slightly so. The type specimen ofGossia diversifolia was (conduplicate) leaf blades. collected from Balade, farther north along the eastern coast Distribution, habitat and phenology. — Known only from from Poindimié, the one known locality of G. conduplicata the type collection at or near Poindimié (Fig. 6) in humid forests sp. nov. Further study is needed to better understand the at 50 meters. Flowering unknown; fruiting in April. differences of these two taxa.

152 ADANSONIA, sér. 3 • 2020 • 42 (7) Revision of New Caledonian Gossia N. Snow & Guymer

A D

F G

Fig. 12. — A-D, Gossia colnettiana (Guillaumin) N. Snow: A, branch and inflorescence; B, bracteoles and flower bud; C, flower minus petals and stamens; D, longitudinal section through flower showing bilocular ovary with axile placentation;E -G, Gossia conduplicata N. Snow, sp. nov.: E, branch with inflorescences;F , leaves (lateral (left) and dorsal (right) views); G, immature fruit views, apical (left) and lateral (right); H-K, Gossia conspicua (Vieill. ex Guillaumin) N. Snow, comb. nov.: H, branch with dense leaves and short internodes; I, strongly recurved leaf blades; J, flower (apical view minus two petals at left), lateral view right( ); K, immature fruit. Vouchers: A-C, MacKee 19108 (BISH); D, Jaffré 3228 (NOU); E-G, MacKee 17222 (NOU); H, I, K, Veillon 5958 (NOU); J, MacKee 5328 (P). Scale bars: A, E, 2 cm; B-D, G, J, K, 2 mm; F, 1 cm; H, I, 5 mm.

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Gossia conspicua (Vieill. ex Guillaumin) midnerve above sulcate proximally becoming flush distally, N. Snow, comb. nov. the secondary (and less frequently) tertiary veins somewhat (Figs 12H-K; 14) raised, oil glands of lower surface present, somewhat sparse and obscure. Myrtus conspicua Vieill. ex Guillaumin, Bulletin de la Société botanique de France 85: 631 (1939). — Austromyrtus conspicua (Guil- laumin) Burret, Notizblatt des Botanischen Gartens und Museums zu Inflorescence Berlin-Dahlem 15: 505 (1941). — Gossia alaternoides (Brongn. & Less than 1 cm (including flower), pedicels 3.5-6.0 mm, Gris) var. conspicua (Vieill. ex Guillaumin) N. Snow, Austrobaileya monads axillary, extrafloral bracts absent. 8: 180 (2010). — Typus: New Caledonia. Grande Terre, South Prov., Montagne de Ouatendé près Gatope, E. Vieillard 2618 (holo-, Bracteoles P[P00602574]; iso-, G[G00341336, image seen], GH[GH00255460], P[P00462848, P00462849]). 2.6-4.0 mm, linear, soon deciduous. Representative herbarium specimen. — P00602574. Hypanthium Etymology. — Derivation uncertain. 3-4 mm, obconic, smooth, lanate; ovary apex indumentum unknown. Distribution, habitat and phenology. — Southeast on Grande Terre (Fig. 14) in maquis over ultramafics,c. 800-1350 m, and from Calyx the northwest at Oua Tilou and Point d’As; flowering December to May; fruiting confirmed January through November. Lobes 5, 1.6-2.1 mm, broadly triangular to narrowly oblong, lanate. Petals and stamens not seen. Specimens examined. — New Caledonia. North Prov., Vallée supérieure de la Riv. Voh, 24.IV.1951, Baumann-Bodenheim 12105 Style (P[P00462818], US, Z); Versant N de l’Oua Tilou, 14.IV.1951, Guillaumin & Baumann-Bodenheim 12361 (P[P00462819], RSA, c. 3 mm (material scant), hairy proximally. US, Z); Slopes of Mt Koniambo, 400-800 m, 31.III.1956, MacKee 4264 (P[P00462839]); Mt. Koniambo, 400 m, 5.I.1961, MacKee Fruits (material scant) 7965 (CANB, P[P00462840], US); Crête Sud du Mont Taom, 5-6 mm, globose, base rounded, green becoming pale purple 1000 m, 31.VII.1981, MacKee 39391 (NOU[NOU028914], to blackish; seeds (only 1 seen) 1 per fruit, c. 3.4 mm. P[P00462842]). — South Prov., Sommet W du Mt. Humboldt, 1400 m, 21.IX.1951, Baumann-Bodenheim 15430 (A, BRI, P[P00459521], US, Z); Mont Oua tendé, 1868, without collector, Remarks (P[P00462846]); Crête de la Montagne des Sources, 8.III.1951, The size, shape, texture, strong marginal recurvation, and Hürlimann 1002 (A, P[P00462824], US, Z); Sommet Point d’As, emergent indumentum of the young leaves of Gossia conspicua 19.XI.1971, Jaffré 535 (NOU[NOU028908], P[P00462830]); comb. nov. most closely resemble Gossia virotii (which see, for Crête de la Montagne des Sources, 1000 m, 13.III.1951, MacKee 2237 (P[P00462836]); Plateau below Montagne des Sources, details). Although subtle, the nodes can be somewhat swollen, 800 m, 23.IX.1956, MacKee 5328 (L, P[P05221714]); Mont thus resembling small platforms from which the petioles arise Kouakoué, south of summit, forest above creek, 21°57’46”S, (e.g., Tronchet 604). One specimen with atypically narrow 166°32’06”E, 1200 m, 9.V.2006, McPherson 19407 et al. (KS- leaf blades (MacKee 39391) resembles Gossia virotii. P[KSP004706], MO, NOU[NOU017142], P[P06668975]); Mont Kouakoué, 1250-1350 m, 30.IV.2006, Pillon 396 et al. (MO[MO-04860792], NOU[NOU012051], P[P06668688]); Mont Kouakoué, 21°58’1”S, 166°32’34”E, 1044 m, 12.I.2002, Gossia diversifolia (Brongn. & Gris) N. Snow Tronchet 604 (MO[MO-04796930], NOU[NOU004501], (Figs 2; 6; 13A-D) P[P00354719]). Austrobaileya 8: 180 (2010). — Eugenia diversifolia Brongn. & Bulletin de la Société botanique de France Description Gris, 12: 180 (1865). — Myrtus diversifolia (Brongn. & Gris) Guillaumin, Bulletin de la Société Shrubs botanique de France 85: 631 (1938). — Typus: New Caledonia. 0.2-1.5 m. Grande Terre, North Prov. Circa Balade, Vieillard 476 (lectotype designated by Snow & Veldkamp 2010: 180); second-stage lecto- Internodes type, here designated, Vieillard 476 (P[P00462912]): epitype, here Mostly < 5 mm near tips of branchlets. designated, Pancher s.n. (P[P00462909], and see comments below). Myrtus flavidaSchltr., Botanische Jahrbucher für Systematik 40 (3): Branchlets Beibl. 92: 30 (1908), syn. nov. — Typus: New Caledonia. South Terete, shortly tomentose on emergence but becoming in- Prov., Le Rat 2003 (type-, B? n.v.). creasingly glabrous. Etymology. — Derived from the Latinization of “diversely leaved” by the authors in the protologue. Leaves Stiffly coriaceous, 2-4 per node; petioles 1.5-3.1 mm, sul- Distribution, habitat and phenology. — Eastern coast of Grande Terre and locally common around Baie de Tina (Fig. 6); in sclerophyl- cate above throughout; blades (1.5-)2.0-3.1 cm, broadly lous forests among hills, sometimes behind mangroves, on black clays linear to narrowly elliptic or narrowly obovate, base cune- or soils derived from schists or calcareous schists; 5-100 m. Flower- ate, margin strongly recurved throughout, apex retuse, ing December through May; fruiting December through October.

154 ADANSONIA, sér. 3 • 2020 • 42 (7) Revision of New Caledonian Gossia N. Snow & Guymer

D B

Fig. 13. — A-D, Gossia diversifolia N. Snow, sp. nov.: A, branch with flowers; B, detail of branchlet with quadrangular-winged young stems; C, flower in bud; D, immature fruit; E-I, Gossia aphthosa (Vieill. ex Brongn. & Gris) N. Snow subsp. longipedunculata N. Snow & Munzinger, subsp. nov.: E, leaves and terminal inflorescence;F , flower detail prior to anthesis;G , apical view of flower with staminal disk (petals and stamens removed); H, branch with leaves and inflorescence; I, maturing fruit. J, K, Gossia kaalaensis N. Snow, sp. nov.: J, branch with inflorescences, petiole (adaxial view) to right, raised adaxial leaf venation near margin (enlarged at upper left); K, immature fruit. Vouchers: A-C, MacKee 43897 (NOU); D, Jaffré 3228 (NOU); E-G, MacKee 26628 (NOU); H-I, MacKee 25749 (NOU); J, K, McPherson 2551 (P). Scale bars: A, J, 2 cm; B, 1 cm; C, D, K, 3 mm; E-G, I, 5 mm; H, 2 mm.

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165°E 166° N

Mt. Panié Ouvéa (1629 m) Island 21°S 21°S Koné

Mt. Humboldt (1618 m) Gossia conspicua (Guillaumin ex Vieill.) N. Snow, comb. nov. Gossia katepahiensis N. Snow, sp. nov. 22° 22° Gossia kuakuensis (Baker f.) N. Snow

100 km Nouméa

164°E 165° 166° 167°

Fig. 14. — Distribution maps of Gossia conspicua (Guillaumin ex Vieill.) N. Snow, comb. nov., G. katepahiensis N. Snow, sp. nov., and G. kuakuensis (Baker f.) N. Snow.

Specimens examined. — New Caledonia. North Prov., Pouem- MacKee 12412, Montravel, parc forestier, 50 m, 9.IV.1965 bout, conservatoire botanique de Tiéa, 5.IX.2000, Dagostini (MO[MO-5813360], NOU[NOU028891]); Nouméa, Baie de 209 (P[P00316468]); Pouembout, 16.II.1972, MacKee 25012 Tina, 16.III.1972, MacKee 25151 (NOU[NOU028890]); Baie de (NOU[NOU028889]); Pouembout, 16.IV.1981, MacK- Tina, 24.III.1974, MacKee 28365 (NOU[NOU028895]); Nouméa: ee 38948 (NOU[NOU028896]); Creek hervouet (propriété Baie de Tina, 3.X.1974, MacKee 29369 (NOU[NOU028898], Johnson), 10.X.2004, Munzinger 1866 (MO[MO-6175968], P[P02089820]); Nouméa: Baie Tina, 4.III.1975, MacKee 29834 NOU[NOU006339], P[P06668680]); Poya, forêt de Nekoro, (NOU[NOU028901]); Nouméa: Baie de Tina, 20.II.1976, Mac- 28.IX.1988, Veillon 6896 (NOU[NOU028902]); Poya, forêt de Kee 30715 (NOU[NOU02899]); Rivage SE Baie La Concep- Neoni, 29.VI.1989, Veillon 7054 (NOU[NOU028897]); Poya, tion, 2.IV.1981, MacKee 38895 (NOU[NOU028900]); Baie propriété Johnston, 10-20 m, 6.V.1998, Veillon 8096 (BISH, de Tina, 12.II.1988, MacKee 43897 (BRI, MO[MO-6751078], NOU[NOU028904], P[P00467803]). — South Prov., Nouméa, 1868- NOU[NOU053243], P[P02089873]); Baie Tina, 10-15 m, 1870, Balansa 126 (A, G, P[P00462889, P00462890, P00462891]); 16.X.1990, Müller 54 (P[P04682695]); Montravel Jardin Bota- Baie de Prony, Herbarium L. Roterau s.n. (P[P00500668]); nique, 19.VI.1965, Schmid 351 (NOU[NOU028907]); Bois de Baie de Tina, 18.V.1985, Hoff 905 (NOU[NOU028906]); Port Desfontes (Nouméa), X.1942, Virot 794 (P[P00500654]); Baie de Tina, 1.VI.1974, Jaffré 1303 (NOU[NOU028894], Bois de Port Desfointes (Nouméa), 14.XI.1942, Virot 817 (A, P[P00462915, P00462916]); Bois de Port Desfointes, 13.XII.1942, P[P0462898, P0462890]); Baie de Tina, 9.IX.1988, Jaffré 2988 Virot 918 (A, MO[MO-04856944], P[P00462918, P00462919]). — (NOU[NOU028905]); Parc forestier de Nouméa, 11.VII.1992, Jaffré Ab loco. 1868, Balansa 416a (Z[Z-000050493]); Balansa 3400 3167 (NOU[NOU028893]); Nouméa, parc forestier, 19.III.1993, (P[P00462894, P00462893]); Baudouin s.n. (P[P00462895]); Jaffré 3228 (NOU[NOU053230]); Mt. Dzumac, 9.X.1909, Le Rat Deplanche 211 (P[P00462897]); Pancher 50 (P[P00462925]); 547 (P[P00462900, P00462901]); Forêt de Magenta, Le Rat 2085 Herbarium Pancher s.n. (G[G00340971, G00340966]); Vieillard (A, P[P004629802]); Route de la prise d’eau, Dumbéa, IX.1904, 476 (P[P00462913]). Le Rat 2269 (P[P00462903]); Port Despointes, 17.IV.1956, Mac- Kee 4420 (A, L, P[P00462904]); Montravel, parc forestier, 30 m, 2.III.1965, MacKee 12178 (NOU[NOU028892], P[P00462905]); Representative herbarium specimen. — P00462909.

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ABC

Fig. 15. — A-C, Gossia katepahiensis N. Snow, sp. nov.: A, branch with inflorescences;B , flowers in bud;C , detail of flower bud (in longitudinal section at right); D-H, Gossia kuakuensis (Baker f.) N. Snow: D, flowering branch; E, detail of simple cyme (note small leafy bracts at base); F, detail of flower (minus petals and stamens); G, young fruit and leaf; H, older branch with attached leaf showing dark, more or less rectangular cracking in epidermal layers; I, J, Gossia mandjeliaensis N. Snow, sp. nov.: I, flowering branch (detail of cracking outer epidermal layers (lower left) and abaxial oil glands (upper left)); J, flower from above (lacking petals and stamens) (left) and laterally (right). Vouchers: A-C, McPherson 5690 (WELTU – from 2 sheets); D-F, Compton 930 (BM); G, Vieillard 2171 (P); H, Vieillard 2218 (P); I-J, McPherson 2551 (P). Scale bars: A, D, 2 cm; B, E-G, 5 mm; C, J, 3 mm; I, 2 mm. Scale bar lacking: H, leaf c. 5 cm long.

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Description time of lectotypification), a second-stage lectotypification is Erect shrubs or treelets provided above. Moreover, epitypification (Art. 9.9,Turland 1.5-4 m. et al. 2018) is desirable given the sterile condition of the lectotype, and that Vieillard 462 at P consists of two sheets, Branchlets both of which lack fertile material. At the present time, the 4-angled; internodes mostly > 5 mm. two sheets of Pancher at P are databased as “types”, but no such indication or annotation appears on the specimens, Leaves and it is unclear at this point whether I was aware of their Coriaceous, 2 per node; petioles 0.8-1.7 mm, round to flattened type status when I first annotated one of these specimens above; blades 1.8-3.1 × 0.9-2.1 cm, ovate to widely ovate, base (P00462908) from Eugenia to Gossia in 2004. Among the rounded to cordate, surface flat or slightly irregular, margin two sheets of Pancher, one in Pancher’s hand (P00462908) slightly undulate, apex obtuse to somewhat acute, midnerve indicates the specifics of the collection on the blue label (on above flush, secondary veins sometimes prominent above, the left), whereas the collecting year of 1862 (also in Pancher’s oil glands of lower surface common but faint (typically most hand) is on the label (to the right), however this specimen abundant along margin). also is sterile. The other specimen of Pancher (P00462909) indicates the year 1861 (printed, not written by hand) on Inflorescence the right-hand label, but details of the collection on the blue 0.6-1.5 cm, of monads (mostly) or simple cymes, terminal label to the left also are in Pancher’s hand (and are nearly or axillary, extrafloral bracts (at base of pedicel) scale-like. identical to those of P004652908), and since this specimen has four flowers, it is selected as the epitype. Bracteoles 0.5-0.8 mm, narrowly ovate to ovate, glabrous to sparsely sericeous. Gossia kaalaensis N. Snow, sp. nov. (Figs 13J-K; 17) Hypanthium 1.4-2 × 2.0-2.5 mm, campanulate, surface smooth (or mi- Petioles narrowly and deeply sulcate, but broadening distally. Leaves narrowly ovate to ovate, margin somewhat undulate, secondary and nutely ribbed in fruit), glabrous; ovary apex shortly hairy. tertiary veins of both surfaces prominently raised. Sepals densely short-sericeous below. Calyx Lobes 5, 0.8-1.8 mm, broadly rounded, glabrous above and Typus. — New Caledonia. Grande Terre, North Prov., Slopes of Mt Kaala, above village, 20°37’39.00”S, 164°23’12.12”S, below, green. c. 550 m, 7.X.1982, G. McPherson 4973 (holo-, P[P00758090]; iso-, BISH[fragment]; MO[MO-3211396]; NOU[NOU030910]). Petals c. 3.5 × 3-4 mm, glabrous above and below, white. Paratypi. — New Caledonia. North Prov. Mont Kaala, 4.IV.1968, Bernardi 12528 (BISH[fragment], L, P[P00462920], US, Z); Voh. Massif Katépaï, au bord d’une ancienne piste, 20°55’49.72”S, Filaments 164°42’02.92”E, c. 470 m, 19.IX.2016, Munzinger 7853 (leg. 2-4 mm, white; anther sacs c. 0.5 mm, pale yellowish. Scoptera) (MPU[640685], P[P02090976]). — South Prov. Kongouaou (= Koungouhaou), 650 m, 2.V.1976, Jaffré 1725 Style (NOU[NOU029644]); Contrefort NW Mt. Taom, 10.V.1979, MacKee 36887 (NOU[NOU029643], P[P02089830]); Crête du c. 3.5 mm, sparsely sericeous below. Oua-Tilou, 20°50’08.99”S, 164°51’27.00”E, 850 m, 18.V.1983, Morat 7398 (NOU[NOU029645], P[P00462921]); Montag- Fruits nes de Balade, Vieillard 484 (left branchlet on mixed collection; 5-8 × 5-7 mm, globular, base rounded, green turning reddish P[P00463007, P00602546]); Oua Tendé près Gatope, Vieillard to black; seeds 2. 2610 (K[K001000439], P[P00462922, P00462923, P00462924]); circ. Kaala-Gomen; S slopes of Mt. Kaala, c. 500 m, 11.VIII.1968, Webster & Hildreth 14719 (DAV, GH, NSW, P[P00462928]). Remarks The combination of 4-angled branchlets, cordate or rounded Representative herbarium specimen. — P00758090. leaf bases, and the tapered fruit base easily diagnose the spe- Etymology. — After the type locality. cies vegetatively from congenerics. The tapered fruit bases can also occur in G. vieillardii. The images of living material Distribution, habitat and phenology. — Grande Terre from were taken in a semi-manicured area (with mowed lawn) in the northwest, where it is most common, sporadically south to Koungouhaou (Fig. 17); in maquis and scrub that grades into low- the Parc Forestier de Nouméa. stature but dense forests, over ultramafics or graywackes, 470-850 m. Among the collections Brongniart & Gris (1865: 180) Flowering April through July; fruiting May through October. mentioned in the protologue (Pancher s.n., [collected in] 1862; Vieillard 462), Snow (Snow & Veldkamp 2010: 182) Description designated Vieillard 462 as the lectotype. Since Vieillard Shrubs 462 consists of two sheets (unknown to the author at the 0.7-2 m.

158 ADANSONIA, sér. 3 • 2020 • 42 (7) Revision of New Caledonian Gossia N. Snow & Guymer

A B

C D

Fig. 16. — A-C, Gossia kuakuensis (Baker f.) N. Snow vel aff.: A, inflorescences and leaves;B , leaves and branchlets with dark rectangular flaking bark;C , young fruits; D, Gossia pancheri. Vouchers: A-C, Fleurot 226 (NOU, P); photo ©Dominique Fleurot, with permission; from Näräjaa, Ponérihouen (http://endemia.nc); D, J.-N. Labat 3486 (P).

Bark Hypanthium Marbled with light and dark gray. c. 2-3 × c. 2.5 mm, campanulate to obconic, surface smooth, glabrous (or sparsely sericeous); ovary apex glabrous. Branchlets Terete to laterally compressed; internodes > 5 mm. Calyx Lobes 5, 1.3-2 mm, broadly ovate to broadly rounded, seri- Leaves ceous-tomentose above, glabrous to sparsely and very short Coriaceous, 2 per node; petioles 3-5 mm, deeply sulcate; sericeous below, drying whitish. blades (4.5-)8.0-9.5 × 1.8-3.4 cm, narrowly elliptic to narrowly ovate or somewhat elliptic, base cuneate (rarely almost Petals rounded), surface flat to somewhat undulate, margin somewhat 3.5-5 × 4-5 mm, sparsely villous above, glabrous below, shortly undulate, apex acute, midnerve sulcate above, secondary veins ciliate on edges. prominently raised above, oil glands of lower surface common, indistinct to prominent (and relatively large or small). Filaments Inflorescence 2.5-4 mm; anther sacs 0.5-0.7 mm. 1.5-3.5 cm, of monads, 3-flowered cymes, or few-flowered racemes, axillary, terminal or sometimes arising from internodes, Style pedicels 0.3-1.3 cm; extrafloral bracts absent or scale-like. 4-5 mm, sparsely sericeous.

Bracteoles Fruits 1-1.5 mm, linear to very narrowly obovate, glabrous to c. 5-5 × 6-7 mm, subglobular to globular, base rounded; green sparsely sericeous. maturing to dark reddish; seeds 2.

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Remarks visible above and below, oil glands of lower surface faint and Several characters are diagnostic for Gossia kaalaensis sp. moderately common. nov. The leaf blades (in sicco) typically are dark green, narrowly elliptic to narrowly ovate, apically acute, and the Inflorescence upper surface is frequently undulate, glossy and has promi- Of monads or elongated racemes, axillary and terminal; ex- nently raised secondary veins. The petiole is deeply sulcate trafloral bracts absent or scale-like. but flattens and broadens distally. The hypanthium often has relatively large, pustular oil glands interspersed among Bracteoles much smaller glands. It most closely resembles Gossia vieil- 1-2 mm, narrowly obovate, glabrous. lardii, but leaves of that species typically are a lighter shade of green. Gossia kaalaensis sp. nov. may also resemble G. Hypanthium pancheri, but the latter has a sericeous hypanthium. 2.5-3 × 2-3 mm, obconic, surface smooth, glabrous to sparsely A specimen with a similar leaf shape and texture to G. kaal- sericeous; ovary apex glabrous. aensis sp. nov. is atypical by the much smaller length of its leaves (Munzinger 7577 [MPU620440]); its placement at Calyx the current time is unresolved. Lobes 5, 1-1.5 mm, broadly rounded, sparsely sericeous near base above, glabrous to sparsely sericeous below, green.

Gossia katepahiensis N. Snow, sp. nov. Petals (Figs 14; 15A-C) 4-5 × 3-4 mm, ciliate on margins, glabrous below.

Resembling Gossia kuakuensis but differing by its narrower bracteoles; Filaments similar to Gossia vieillardii but having rounded leaf bases; and differing fromGossia bourailensis sp. nov. by the broadly obtuse to 3-5 mm; anther sacs 0.4-0.6 mm. rounded apex of the leaf. Style Typus. — New Caledonia. Grande Terre, North Prov., Mt. c. 5 mm, sericeous. Katépahié, above Voh, 20°55’58.08”S, 164°41’33.00”E, 4.V.1983, c. 350 m, McPherson 5690 (holo-, P[P00402743]; iso-, BISH[fragment], BRI, MO[MO-3183836], NOU[NOU030911], Fruits (young) NSW, WELTU[2 sheets]). 7-8 mm, ellipsoid and somewhat tapered at the base, light green but beginning to darken; seeds up to 2. Paratypi. — New Caledonia. North Prov., Voh, Massif Ka- tépahï, en haut d’un sentier de randonnée, 20°55’26.36”S, 164°42’16.94”E, c. 530 m, 19.IX.2016, Munzinger 7867 (leg. Remarks Scoptera) (MPU[MPU640688]; P[P02090939]). This species resembles several others in Gossia. Its leaf apex is rounded to broadly obtuse (and sometimes slightly retuse), and Representative herbarium specimen. — P00402743. the secondary veins are visible in sicco. Mature inflorescences Etymology. — Derived from the type locality. typically are longer than the adjacent leaves, and the inflorescence and pedicels are relatively thin and often somewhat bowed. In Distribution, habitat and phenology. — Known from two collections on Mt. Katépahie (Fig. 14) from forest remnants over addition, the base of the petals is thicker than most other species. ultramafics from approximately 350-530 meters. Flowering in May; A specimen I annotated in 2004 with this name fruiting in August. (P[P00402742]) likely represents a different taxon, given its cuneate leaf bases and shorter inflorescences; its identity is Description uncertain from the digital image. Rounded shrubs to trees A specimen allegedly from Tahiti by an unknown collector 1.5-3 m. (Anonymous, « Commun à Tahiti » ([P05094318])) was identified (but not annotated) by some other person. Although its Bark relatively large inflorescence resembles that of the type speci- Light gray to almost white and longitudinally furrowed. men, its different leaf morphology and the distance between Tahiti and New Caledonia (c. 4700 km) together make such Branchlets an identification unlikely. Terete, bright red on emergence; internodes > 5 mm.

Leaves Gossia kuakuensis (Baker f.) N. Snow Coriaceous, 2 per node; petioles 2.5-4 mm, sometimes slightly (Figs 14; 15D-H; 16A-C) sulcate; blades 2.2-5.5 × 1.5-3.5 cm, elliptic, base rounded, Austrobaileya 8: 181 (2010). — Psidium kuakuense Baker f., surface flat, margin slightly recurved and surface slightly Journal of the Linnaean Society, Botany 45: 318 (1921). — Typus: undulate, apex rounded to broadly obtuse (and sometimes New Caledonia. Grande Terre, South Prov., Kuakué (= Kouakoué), slightly retuse), midnerve sulcate above, secondary veins faintly 13.V.1914, Compton 930 (holo-, BM[BM000581652]).

160 ADANSONIA, sér. 3 • 2020 • 42 (7) Revision of New Caledonian Gossia N. Snow & Guymer

Psidium floribundumVieill. ex Guillaumin, nomen nudum, Annales du Calyx Musée colonial de Mareille, sér. 2, 9: 150 (1911). — See discussion in Lobes 5, 1.5-3 mm, broadly rounded, ciliate but otherwise Snow & Veldkamp (2010: 181) regarding this unpublished name. glabrous, green. Eugenia cataractarum Guillaumin, Bulletin de la Société botanique de France 85: 636 (1939). — Austromyrtus cataracarum (Guil- Petals laumin) Burret, Notizblatt des Botanischen Gartens und Museums 4-6 × 3-5 mm, white, sparsely ciliate on margins. zu Berlin-Dahlem 15: 503 (1941). — Typus: New Caledonia. Grande Terre, South Prov., Secus ripus torentium prope Touho, Vieillard 2171 (lecto-, P[P00602576] designated here; isolecto-, Filaments G, GH[GH00071539], P[P00751830]). 2.5-4.5(-6) mm; anther sacs c. 0.5 mm. Style 3-5(-6.5) mm, glabrous to sparsely sericeous. Specimens examined. — New Caledonia. North Prov., Näräjaa, Ponerihouen, 550 m, 26.III.2016, Fleurot 226 (P[P01168448]); Pente Fruits Sud du Mt. Kaala, 12.VI.1965, MacKee 12758 (NOU[NOU029641], P[P00462879, P00462880]); Pente Sud-Ouest du Mt. Kaala, 5.5-10 × 5.5-10 mm, subglobular to globular, base rounded, 300-600 m, 10.IX.1967, MacKee 17490 (NOU[NOU029640], green when young (mature color unconfirmed); seeds 1-4. P[P02089978, P05094102); Massif de Tion-non, 800-900 m, 21.III.1968, MacKee 18516 (P[P02089982]; Col d’Amieu-Ouambéa, Remarks 500-600 m, 31.III.1970, MacKee 21745 (NOU[NOU030905]); The bark of branchlets and main trunk typically breaks into Bord des torrents, Wagape, Vieillard 2171bis (BISH, P[P00462881, P00462882]); Bord des torrents à “Poinendu”, Vieillard 2218 dark rectangular flakes, and the branches reportedly can be (G[G00340965, G00340992, images seen], MEL n.v., P, 2 sheets). pendant. Some leaves on the holotype are atypically clasping. The peduncles in the inflorescence are glabrous but can be Representative herbarium specimen. — P02089978. prominently striate. We tentatively here include Fleurot 226 (Fig. 16A-C in Etymology. — From the type locality. Näräjaa, Ponérihouen), although its leaves are sessile and Distribution, habitat and phenology. — Scattered on Grande slightly clasping, whereas other specimens have short petioles, Terre (Fig. 14); in maquis on ultramafics or edges of dense moist and its fruits have 4-merous calyx lobes. Study of herbarium forests over sedimentary volcanics, 100-600 m (but more data needed; vouchers (unavailable before submission) will help determine see remarks). Flowering March through June; fruiting confirmed only for September. its proper placement.

Description Gossia mandjeliaensis N. Snow, sp. nov. Shrubs or trees (Figs 15I-J; 17) 1-6 m. The leaves resemble those of Gossia vieillardii, but the oil glands on the lower leaf surface of G. mandjeliaensis sp. nov. are conspicuous Branchlets and dense and the base of its hypanthium somewhat rounded. Terete to compressed (the older branchlets with dark, rectangular flakes); internodes > 5 mm. Typus. — New Caledonia. Grande Terre, North Prov., Forested slopes below radio tower of Mandjélia, c. 5 air-km W of Poue- bo, 20°23’58.99”S, 164°31’30.00”E, 12.IV.1980, McPherson Leaves 2551 (holo-, P[P00402744]; iso-, BISH, MO[MO-3226084], Coriaceous, 2 per node; petioles 1.5-4 mm (or leaves sessile), NOU[NOU030914], WELTU). sometimes slightly sulcate; blades (3-)4.5-9.0 × 3.0-6.0 cm, elliptic to broadly elliptic, base rounded to cordate or clasp- Paratypi. — McPherson 6327, Mandjélia, above Pouébo, 20°24’18.40”S, 164°31’05.00”E, c. 650 m, 21.II.1984 (MO[MO-3225874], ing, surface flat to somewhat creased along midvein, margin NOU[NOU029639], P[P00462929], WELTU). flat to slightly irregular, apex obtuse, midnerve above flush to slightly sulcate at base, secondary veins flush above, oil glands Representative herbarium specimen. — P00402744. of lower surface dense but indistinct. Etymology. — After the type locality. Inflorescence Distribution, habitat and phenology. — Known only from 2.5-8.5 cm, a dichasium, raceme or panicle, terminal or Mt. Mandjélia (Fig. 17); forested slopes over metamorphic rock, 600-650 m. Flowering February through April; fruiting period axillary; pedicels (1.5-) 4-11 mm; extrafloral bracts leaflike. unknown. Bracteoles 1.5-4.5 mm, linear to narrowly elliptic, glabrous to sparsely Description sericeous. Trees To 10 m. Hypanthium Obconic, 2.3-2.6 × (2-)3.4-3.8(-4.2) mm wide, surface smooth, Branchlets glabrous; ovary apex glabrous. Terete; internodes > 5 mm.

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Leaves Typus. — New Caledonia. Grande Terre, South Prov., Île des Pins, Coriaceous, 2 per node; petioles 2-3 mm, sometimes slightly Pic Nga near Kuto, 22°39’18.00”S, 167°27’24.98”E, c. 100 m, sulcate; blades (1.5-)3.0-6.0 × (-1.0)2.0-4.0 cm, narrowly el- 14.VIII.1982, G. McPherson 4878 (holo-, MO[MO-3210881]; iso-, BISH fragment, NOU[NOU052879]). liptic to broadly elliptic, base cuneate (or occasionally rounded), surface flat, margin flat, apex obtuse to acute, midnerve above Representative herbarium specimen. — No images currently available. sulcate near base but becoming flush distally, secondary veins Etymology. — Derived from Pic Nga, the locality of the type specimen. faint and flush above and below, oil glands dense but typically indistinct above and below. Distribution, habitat and phenology. — Known only from the type collection from Pic Nga near Kuto on Île des Pins (Fig. 19); in scrub over Inflorescence ultramafics along a creek,c. 100 m. Flowering August; fruiting August. Up to 3.5 cm, of monads, simple cymes, or few-flowered racemes, axillary or terminal, pedicels 0.1-0.8 cm; extrafloral Description bracts absent or scale-like. Shrub To 2 m. Bracteoles 1.5-3 mm, linear, very narrowly elliptic, or narrowly obovate, Branchlets more or less sericeous. Terete to strongly compressed; internodes > 5 mm.

Hypanthium Leaves 3-4 × 3-4 mm, campanulate, surface smooth, sericeous; ovary Coriaceous, 2 per node; petioles 3-5 mm, sometimes slightly apex sparsely hairy. sulcate; blades (2.8-)5.0-8.3 × 1.2-4.0 cm, elliptic, base cuneate, surface flat, margin flat to slightly revolute, apex broadly Calyx obtuse and sometimes slightly retuse, midnerve sulcate in Lobes 5, 1.3-1.9 mm, broadly rounded, glabrous above, seri- lower half, secondary veins flush to slightly raised above, oil ceous near base below and ciliate on edges, green. glands of lower surface dense and prominent.

Petals Inflorescence 4-5 × 4-5 mm, glabrous above, glabrous to sparsely sericeous Of single flowers or simple cymes, terminal or axillary; ex- below. trafloral bracts absent.

Filaments Bracteoles 3-5 mm; anther sacs 0.5-0.6 mm. Unknown.

Style Hypanthium c. 5 mm, sericeous at base. 4.5-7 × 4-5 mm wide (in fruit), obconic, surface smooth, glabrous; ovary apex glabrous. Fruits Dimensions unknown, said to be green, mature fruit un- Calyx known; seeds not seen. Lobes 5, 1.7-2.5 mm, broadly triangular to broadly rounded, sparsely short ciliate near apex, greenish. Remarks Gossia mandjeliaensis sp. nov. differs from other New Cal- Petals edonian congenerics by subtle traits, including the dense but c. 5 × 5 mm, glabrous above, glabrous to sparsely sericeous below. small oil glands on the leaf surfaces; a glabrous to slightly hairy staminal disk; sericeous indumentum on the pedicels, Filaments abaxial surface of calyx lobes, and style; and faint abaxially 4-6 mm; anther sacs 0.4-0.6 mm. secondary leaf veins. Gossia colnettiana is somewhat similar, but has fewer oil glands abaxially, raised abaxial secondary Style veins, ferrugineous sericeous hairs in flower, and a hairy c. 5 mm, sparsely sericeous. staminal disk. Fruits 5.5-7 × 5-7.5 mm, globular, base rounded, green (immature); Gossia ngaensis N. Snow, sp. nov. seeds 1-2. (Figs 18A-C; 19) Remarks Resembling Gossia nigripes but with longer and flatter leaf blades Gossia ngaensis sp. nov. allegedly was photographed on Pic (margins not recurved), and without an obtuse apex. Nga during September 2017 (G. Gâteblé pers. comm. 2018),

162 ADANSONIA, sér. 3 • 2020 • 42 (7) Revision of New Caledonian Gossia N. Snow & Guymer

165°E 166° N

Mt. Panié Ouvéa (1629 m) Island 21°S 21°S Koné

Gossia kaalaensis N. Snow, sp. nov. Mt. Humboldt (1618 m) Gossia mandjelianensis N. Snow Gossia nigripes (Guillaumin) N. Snow

22° Gossia ouazangouensis N. Snow, sp. nov. 22°

100 km Nouméa

164°E 165° 166° 167°

Fig. 17. — Distribution maps of G. kaalaensis N. Snow, sp. nov., G. mandjeliaensis N. Snow, G. nigripes (Gullaumin) N. Snow, and G. ouazangouensis N. Snow, sp. nov. but no voucher was taken and its identification was not con- Distribution, habitat and phenology. — Mt Mandjélia and firmed. The species most closely resemblesG. nigripes (see areas to the north and from the type collection south of Canala key for differences). It also resemblesG. mandjeliaensis sp. (Fig. 17); over micaschistes and forests, sometimes as a dominant on montane ridges, c. 500-1000 m. Flowering February and March; nov., but its hypanthium is sericeous and it occurs on Massif fruiting June through December. du Panié on the northeast part of Grande Terre. Specimens examined. — New Caledonia. Grande Terre, North Prov. Pouébo, 600 m, 24.VII.1974, MacKee 28968 (P[P02089864]); Mt Gossia nigripes (Guillaumin) N. Snow Mandjélia; 19.XII.1981, McPherson 4509 (BISH[fragment], MO, NOU[NOU029638], NSW, P[P05094545], PTBG, WELTU); (Figs 17; 18D, E) above Ouegoa along forest road, 18.VIII.1981, Mueller-Dombois 81081810 (BISH); ibid. loco, Mueller-Dumbois 81081831 (BISH); Austrobaileya 8: 181 (2010). — Myrtus nigripes Guillaumin, Col d’Amoss, 20°18’22”S, 164°23’51”E, 14.X.2002, Tronchet 364 Bulletin de la Société botanique de France 85: 632 (1939). — Aus- et al. (K n.v., MO, NOU, P[P00354952]); Haut Diahot, Tendé, tromyrtus nigripes (Guillaumin) Burret, Notizblatt des Botanischen 600 m, 30.VI.1982, MacKee 40566 (MO, NOU[NOU029624], Gartens und Museums zu Berlin-Dahlem 15: 505 (1941). — Typus: P[P02089875]). New Caledonia. Grande Terre, South Prov., Montagnes situées au Sud de Canala, 20.XI.1869, B. Balansa 2085 (holo-, P[P00602577]; Representative herbarium specimen. — P02089875. iso-, P[P00462930, P00462931]).

Description Etymology. — Derivation from the Latin niger (black) and pes (foot) in reference to the petioles (foot of the blade), which dry Shrubs or trees dark brown to blackish. 3-8 m.

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B A

E D

Fig. 18. — A-C, Gossia ngaensis N. Snow, sp. nov.: A, adaxial surface of leaf (right) with detail (left) venation; B, tip of branch with solitary young fruit; C, seed (left) and fruit (right). D, E, Gossia nigripes (Guillaumin) N. Snow: D, branch with fruits; E, detail of leaf (with dark petioles) and fruit; F-I, Gossia ouazangouensis N. Snow, sp. nov.: F, flowering branch with abaxial oil gland details lower( left); G, branchlet showing cracking epidermal layer; H, detail of cyme; I, detail of flower prior to anthesis. Vouchers: A-C, McPherson 4878 (NOU); D-E, McPherson 4509 (WELTU); F-I, MacKee 16927 (P). Scale bars: A, B, D, 2 cm; C, E, F-I, 5 mm.

164 ADANSONIA, sér. 3 • 2020 • 42 (7) Revision of New Caledonian Gossia N. Snow & Guymer

164°E 165° 166° 167° 168° 19°S 19°S VANUATU

Gossia ngaensis N. Snow, sp. nov. Gossia vieillardii (Brongn. & Gris) N. Snow 20° Aneityum 20°

Mt. Panié Ouvéa (1629 m) Island

Loyalty Islands 21° Koné 21°

NEW CALEDONIA

Mt. Humboldt (1618 m) 22° 22°

Nouméa Isle of Pines 200 km

164°E 165° 166° 167° 168° 169° 170°

Fig. 19. — Distribution maps of Gossia ngaensis N. Snow, sp. nov. and G. vieillardii (Brongn. & Gris) N. Snow.

Branchlets Calyx Terete; internodes > 5 mm. Lobes 4 (5 occasionally), 1-2 mm, broadly ovate to rounded, glabrous above and below, green. Leaves Coriaceous, 2 per node; petioles 2.5-4 mm, terete; blades 2.0- Petals 4.5(-4.8) × 0.8-1.7(-2.4) cm, narrowly elliptic to narrowly Not seen. obovate, base narrowly cuneate, surface flat, margin flat, apex obtuse, midnerve above sulcate proximally becoming flush Filaments apically, secondary veins not visible above, oil glands of lower 3-4.5 mm; anther sacs 0.6-0.8 mm. surface dense but indistinct. Style Inflorescence 4.5-5 mm, hairy to near apex. Of monads or 3-flowered cymes, axillary, pedicels to 4.5 cm; extrafloral bracts absent. Fruits 5-9 × 6-11 mm, subglobular to globular, base rounded, green becom- Bracteoles ing reddish then bluish-black; seeds 2-3, irregularly hemispherical. c. 2 mm, narrowly ovate to ovate, glabrous. Remarks Hypanthium Within Gossia the species is distinguished by leaves that are 2.8-3.2 × c. 3 mm, urceolate, surface smooth, glabrous; ovary nearly all 4 cm or less long and distinctly yellowish-green, apex glabrous. with dark maroon petioles that dry to nearly black.

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The species can resemble Gossia vieillardii, but that species Bracteoles has longer leaf blades on average, longer inflorescences, and 0.9-1.2 mm, linear to narrowly elliptic, sparsely sericeous. fruits that often taper distinctly at the base. Additional data from flowering material are needed. Hypanthium 2-3 × 2-3.5 mm, obconic, surface smooth, glabrous.

Gossia ouazangouensis N. Snow, sp. nov. Calyx (Figs 17; 18F-I) Lobes 5, 1.1-2.8 mm, broadly rounded, glabrous above but ciliate on margins, glabrous below, drying whitish. The species can be distinguished by its thin whitish calyx lobes, glabrous hypanthium, and a mostly terminal to subterminal infloresence that significantly exceeds the length of the leaves. It somewhat resembles Petals Gossia kaalaensis sp. nov., but the adaxial leaf surface of the latter is 6.0-6.5 × 5-6 mm, glabrous above but ciliate on margins, glossy and has prominently raised secondary veins and an acute apex, glabrous below, whitish. and thicker and greenish calyx lobes. In contrast, the adaxial leaf surface of G. ouazangouensis sp. nov. is matte with flush secondary veins and an obtuse apex and thin, whitish or whitish-green calyx lobes. Filaments 4-5 mm, pinkish to the base; anther sacs c. 0.5 mm. Typus. — New Caledonia. Grande Terre, North Prov., pente Ouest de Mt Ouazangou, 20°44’47.00”S, 164°27’56.16”E, 19.VI.1967, MacKee 16927 (holo-, P[P00751831]; iso-, BISH[fragment], Style NOU[NOU030913]). 5-6 mm, sparsely villous, especially in lower half; ovary apex hairy at base of style. Paratypi. — New Caledonia. Grande Terre. North Prov., Poum, 30 m, 27.IV.2004, G. Dagostini 843 & Nigote (NOU, P[P05121896, P05121897], WELTU n.v.); Mont Ouazangou Pente Ouest, Fruits 20°44’47.00”S, 164°27’56.16”E, 300-500 m, 4.III.1972, MacKee Unknown. 25123 (NOU[NOU029637], P[P02089986] with dups [n.v.] to distribute to BISH, CHR, G, MO, NSW, RB, TNS]); Kaala-Gomen, Remarks Massif Onajiele, 20°46’07.02”S, 164°27’44.65”E, c. 100 m, 16.V.2015, Munzinger 7592 (leg. Scoptera) (MPU[MPU620439]). — South The inflorescence branches on MacKee 25123 are somewhat Prov., Vallée de la Tontouta, 21°56’36.99”S, 166°17’35.99”E, 0-50 m, shorter and more gracile, and the branches more reddish and 22.III.2006, Pillon 329 et al. (NOU[NOU011598], P[P06668689]). with a flakier epidermis, but the leaf morphology, locality and Representative herbarium specimen. — P00751831. substrate of the two collections otherwise match well. An up- laced 4-merous specimen (Munzinger 7890 [MPU640674; Etymology. — From the type locality. P020290978]) may be this species. Distribution, habitat and phenology. — Known from the western peak of Mont Ouazangou (Massif Onajile) and Poum in the northwest, and Tontouta in the southwest (Fig. 17); in maquis on Gossia pancheri (Brongn. & Gris) N. Snow ultramafics from near sea level to 500 m. Flowering approximately April through June; fruiting unknown. (Figs 16D; 20; 21A-D)

Description Austrobaileya 8: 181 (2010). — Eugenia pancheri Brongn. & Gris, Bulletin de la Société botanique de France 12: 180 (1865). — Austro- Tree-like shrubs myrtus pancheri (Brongn. & Gris) Burret, Notizblatt des Botanischen To 2-5 m. Gartens und Museums zu Berlin-Dahlem 15: 503 (1941). — Typus: New Caledonia. Grande Terre, South Prov., Montagnes de Yaté, Vieillard Bark 508 (lecto- [designated by Snow & Veldkamp 2010], P[P00462968]; Light reddish or grayish and craking longitudinally. isolecto-, P[P00462969, P00469970, P00602555]). — Snow annotated the lectotype in 2004 but did not specify a barcode number (Snow & Veldkamp 2010: 181), which at that time did not exist. Branchlets One of the isolectotypes (P00602555) was annotated by the author in Terete; internodes > 5 mm. 2005 mistakenly as “lectotype”, instead (correctly) of “isolectotype”. Myrtus luteo-viridis Baker f., The Journal of the Linnean Society, Leaves Botany 45: 312 (1921). — Austromyrtus luteo-viridis (Baker f.) Coriaceous, 2 per node; petioles 1.5-3 mm, sulcate; blades Burret, Notizblatt des Botanischen Gartens und Museums zu Ber- (2.5-)6.0-7.5 × 1.2-3.3 cm, ovate, base cuneate, surface flat, lin-Dahlem 15: 505 (1941). — Typus: New Caledonia. Grande margin slightly revolute throughout, apex obtuse, midnerve Terre, South Prov., Ngoye Mts. to NW, Compton 1339 (holo-, sulcate in lower half, secondary veins mostly flush above, oil BM[BM000581650], photo at BISH]). glands of lower surface common but faint. Eugenia angustibracteolata Baker f., The Journal of the Linnean Soci- ety, Botany 45: 313 (1921). — Typus: New Caledonia. Grande Inflorescence Terre, South Prov., Presqu’île Bogota, 13.V.1914, Compton 1002 1.5-4.5 cm, mostly of 3-flowered cymes (infrequently of (holo-, BM[BM000581654], photo at BISH). monads), axillary or terminal; extrafloral bracts absent or Specimens ecamined. — New Caledonia. North Prov., Boulinda près scale-like. campement, 500 m, 23.IV.1968, Jaffré 18 (NOU, P); Massif du Ko-

166 ADANSONIA, sér. 3 • 2020 • 42 (7) Revision of New Caledonian Gossia N. Snow & Guymer

niambo, 250 m, 18.XI.1971, Jaffré 516 (NOU[NOU029667]); Massif McPherson 4451 (MO, NOU[NOU029656], NSW, WELTU); by du Koniambo, 900 m, 20.XII.1972, Jaffré 1038 (NOU[NOU29662]); Fausse Yaté River, 23.XI.1982, McPherson 5185 (BISH, MO); Forested Kouaoua: Mea, 12.I.1978, Jaffré 2275 (NOU[NOU029649]); Mont slopes above a tributary of the Rivière Ni, 750-770 m, 12.XI.2003, Boulinda, pente au-dessus de la Oua-Népoua, 750-850 m, 26.VII.1967, McPherson 19230 (BISH, MO, NOU); Me Adeo piste en direction du MacKee 17177 (NOU[NOU029683]); Taom, Mt. Homédéboa, 800- Menazi, 24.III.1988, Morat 8008 (NOU[NOU029655]); Sommet du 900 m, X.1969, MacKee 20995 (NOU[NOU029680]); Mont Do, Col de Yaté, 26.VII.1965, Schmid 506 (NOU[NOU029672]); Mine 950-1020 m, X.1969, MacKee 21089 (NOU[NOU029677); Ma- Pauline, vers 450 m, 18.III.1968, Veillon 1648 (NOU[NOU029673]); nambo, 500 m, 29.IX.1989, MacKee 44623 (NOU[NOU0029652], Route vers Prony, après Col de Crève-Coeur, 14.X.1968, Veillon 1859 P[P02089832]); Poro, Kaéoua, 500 m, 17.XII.1992, MacKee 46074 (NOU[NOU029674], P[P00462967, P05094569], WELTU). — Ab (P[P02089865]); Contrefort Nord-Ouest du Mt. Taom, 5.X.1979, loco. Deplanche 530, 1861 (P); Deplanche 531, 1861 (P); Deplanche MacKee 36887 (P); Plateau de Ouazangou, 26.III.1980, MacKee 64172, 1861 (P). 37951 (P[P02089833]); Mont Taom (Crête Ouest), 900 m, 8.I.1981, MacKee 38519 (NOU[NOU029650], P[P02089884]); Mont Taom, Representative herbarium specimen. — P02089884. 900 m, 31.VII.1981, MacKee 39418 (MO, NOU[NOU029647], P[P02089883]); Poro, Kaseoua, 500 m, 17.XII.1992, MacKee 46074 Etymology. — Named for Jean Armand Isidore Pancher (1814‒1877), (NOU[NOU029653], P[P02089865]); Mt Koniambo, c. 700 m, French botanist who worked in New Caledonia. 13.X.1982, McPherson 5019 (MO, NOU[NOU053242]); Mt Ko- niambo, c. 550 m, 5.I.1983, McPherson 5297 (BRI, MO, NOU[- Distribution, habitat and phenology. — Widespread on Grande NOU029654], NSW, PTBG, WELTU); Mt. Colnett, 800-925 m, Terre (Fig. 20); in maquis, forsest edges, and humid forests, often on 2.XI.2003, McPherson 19142 (BISH, MO, NOU[NOU004366], P); ultramafics, 150-1300 m. Flowering throughout the year; fruiting Est du Kopeto, 900 m, 28.XI.1980, Morat 6642 (NOU[NOU029671]); mostly December through July. Massif du Tchingou, face est, 950 m, 29.III.2001, Munzinger 563 (MO, NOU[NOU029668]); Massif du Tchingou, face est, 1000 m, 31.III.2001, Munzinger 588 (MO); Massif du Tchingou, face est, Description 1120 m, 31.III.2001, Munzinger 615 (MO, P); Paéoua Mt, 800-1100 m, Shrubs or trees 26.X.2010, Munzinger 6017 (MO, P). — South Prov., Pente brûlée 1.5-10 m. près de la mine du “Mois de Mai”, 300 m, 13.VIII.1951, Baumann- Bodenheim 15115 (A, US, Z); Canala, Presqu’île de Bogota, c. 450 m, 3.VII.1997, Dagostini 120 (NOU[NOU029675]); Goro-Nickel, rel- Branchlets evé 34, 20.VI.2002, Dagostini & Rigault 505 (NOU[NOU029679]); Terete to compressed, surface developing small dark rectan- Goro, concession SLN “Camille”, 18.V.2002, Dagostini 615 (MO, gular flakes; internodes > 5 mm. NOU[NOU029678]); Col de Mouirange, forêt Desmazure, 10.III.2005, Dagostini 1050 (NOU[NOU009280]); Montagne des Sources, Leaves 3.XII.1992, Dawson WELTU 16573 (NOU[NOU029676], WELTU); Bord de la Fausse Yaté, 10.VII.1981, Hoff 3581 (NOU[NOU029659]); Coriaceous, 2 per node; petioles 2-6.5 mm, sulcate; blades Dans la partie supérieure de la vallée de l’Odjijoni, 180 m, 2.VI.1951, 4.0-8.5 × 1.7-3.8 cm, elliptic, base cuneate, surface flat, mar- Hürlimann 1463 (US, Z); Vallée latérale de la Pourina, 160 m, 4.VI.1951, gin slightly revolute, apex obtuse or retuse, midnerve above Hürlimann 1482 (US, Z); au-dessus du campement Bernier (Mon- sulcate, secondary veins above raised prominently, oil glands tagne des Sources), 850 m, 13.XI.1951, Hürlimann 3019 (US, Z); of lower surface dense (and of uniform size), prominent. Plaine des Lacs, 15.IV.1969, Jaffré 18 (NOU[NOU029660]); Plaine des Lacs, 15.IV.1969, Jaffré 200 (NOU[NOU029658]); Région de Kouaoua, 30.IX.1969, Jaffré 275(NOU[NOU29661]); Kouaoua, Inflorescence 30.IX.1969, Jaffré 294 (NOU[NOU029664]); Mt. Kouakoué, 2-5.5(-8) cm, pedicels (1.5-)5-9 mm, a simple cyme, raceme, 21°58’43”S, 166°32’15”E, 980-1050 m, 9.X.2004, Labat 3486 (K or panicle, terminal or axillary; extrafloral bracts leafy. n.v., MO, NOU[NOU009917], P[P00454747], WELTU); 12 km S of Riv. des Pirogues, ridge overlooking Prony, 200 m, 15.X.1955, Bracteoles MacKee 3247 (US); Yaté road near barrage, 150-200 m, 6.V.1956, MacKee 4494 (L); Route de la Montagne des Sources, 500-700 m, 1.3-2 mm, linear to narrowly elliptic, sparsely to densely sericeous. 27.X.1956, MacKee 5665 (L); Vallée de Mamié, 200 m, 6.X.1966, MacKee 15740 (NOU[NOU029681]); Ouroué, 21.VII.1967, MacKee Hypanthium 17148 (P[P02089831]); Rivière du Humboldt, 800-900 m, 29.X.1967, c. 3 mm × 3-3.5 mm, campanulate, surface smooth, sericeous MacKee 17838 (NOU[NOU029648]); Thio-Mt Nékandi, 1200- to densely sericeous; ovary apex densely tomentose. 1300 m, 7.XI.1967, MacKee 17925 (NOU[NOUO29862]); Mé Ori, Plateau sommital, 900-1000 m, 3.XII.1969, MacKee 21205 (NOU[NOU029657]); Vallée de la Ouinnée, 25.V.1974, MacKee Calyx 28691 (NOU[NOU029666]); Yaté, plateau au Sud du village, Lobes 5, 1.3-2.5 mm, broadly rounded to almost truncate, 300 m, 4.VI.1977, MacKee 33235 (NOU[NOU029665]); Dalma- sericeous to densely sericeous above and below, green. tes Les (Mouirange Col), 150 m, 29.II.1980, MacKee 37865 (NOU) [NOU029608]); Vallée de la Ni, 100 m, 3.XII.1981, MacKee 40035 (NOU[NOU029651]); Hauteurs de Yaté, 250 m, 10.III.1985, Petals MacKee 42493 (MO, NOU[NOU029669], P[P02089882]); Along 4.5-6 × c. 5 mm, somewhat tomentose above and below. old lumber road to top of Mt Mé Ori, above Katrikoin, 8.IX.1980, McPherson 3069 (BRI, MO, NOU[NOU029646], NSW, WELTU); Filaments Montagne des Sources, c. 700 m, 25.X.1980, McPherson 3275 (BRI, 2.5-4 mm; anther sacs 0.5-0.8 mm. MO, NOU[NOU029663], NSW, WELTU); Along Nouméa-Ya- té road, near descent towards Yaté, 400 m, 8.VII.1981, McPherson 3894 (MO, NOU[NOU20670], WELTU); Upper reaches of valley Style Rivière des Pirogues, N of Nouméa-Yaté road, c. 300 m, 12.XII.1981, c. 4 mm, lower half villous.

ADANSONIA, sér. 3 • 2020 • 42 (7) 167 Snow N.

Fruits (material scant) Representative herbarium specimen. — P00462973. 5-6 × 5-6 mm, dark blue-black, globose, base rounded; seeds 1-2. Etymology. — From the Latin ramus (branch) and flos(flower), in reference to the flowers mostly arising from naked branches (before Remarks new leaves appear on young branches or proximal along branches Gossia pancheri is the second most common member of the genus to existing leaves). in New Caledonia after Gossia vieillardii. It is common in the Distribution, habitat and phenology. — Central Grande Terre, southern part of Grande Terre but ranges north to Mt. Colnett. especially in the area of Col d’Amieu (Fig. 20); in humid forests over The best distinguishing characters include the raised second- ultramafics and schistes, 150-700 m. Flowering October through ary and tertiary venation of the leaves, the thickly coriaceous April; fruiting November through September. texture and often glossy sheen of the upper leaf surface. The species most closely resembles Gossia kaalaensis sp. nov., Description but that species has narrower leaves, a more acute leaf apex, Shrubs and trees and a glabrous ovary apex. Some specimens (e.g., MacKee 3.5-10 m. 28691 and 42493) have more densely pubescent and somewhat smaller flowers, but these differences are merely quantitative. Branchlets Terete; internodes > 5 mm.

Gossia ramiflora N. Snow, sp. nov. Leaves (Figs 20; 21E-I) Coriaceous, 2 per node; petioles (1.5-)3-6 mm, sometimes slightly sulcate; blades (4.5-)6.5-10.5 × (-2.0)3.0-5.0(-6.0) Diagnosed by others in the genus by the prominently undulating cm, elliptic to occasionally ovate, base mostly cuneate to leaf margins and (mostly) ramiflorous inflorescences. occasionally rounded, surface undulate, margin moderately Typus. — New Caledonia. Grande Terre, South Prov., Farino: Forêt to strongly and irregularly undulate throughout, apex acute, Mépéou, [21°37’5.016”S, 165°46’35.004”E], 500 m, 22.II.1966, midnerve above sulcate, secondary veins flush above and MacKee 14417 (holo-, P[P00462973]; iso-, CANB, K n.v., L n.v., below, oil glands of lower surface dense and prominent. MO[MO2907912], NOU[NOU029712], P[P00462974], WELTU).

Paratypi. — New Caledonia. Grande Terre, North Prov., Route Inflorescence Tiwaka Kono, 150 m, 30.I.1987, Jaffré 2844 (NOU[NOU029710], 1.0-4.5 cm, pedicels 0.2-0.8 cm, racemes or panicles, mostly P[P00462983]); Western slope of Plateau de Dogny, 600-900 m, naked on branches proximal to leaves or axillary, extrafloral 10.XI.1958, MacKee 6537 (P[P00462984]); Along trail from bracts absent or scale-like. Sarraméa toward Plateau de Dogny, c. 450 m, 1.IX.1982, McPher- son 4924 (BISH[fragment], BRI, MO[MO-3211402], NOU[- NOU029702], P[P00462975]. — South Prov., Col d’Amieu, Bracteoles 500-600 m, 13.VII.1965, Bernardi 9611 (NOU[NOU029711], 1-2 mm, narrowly elliptic or (less so) narrowly triangular, P[P00462982]); Col d’Amieu, Vallé de Toili, 350-400 m, sparsely sericeous. 19.I.1965, MacKee 12002 (NOU[NOU029705], WELTU); Farino, Forêt du Pic Noir, 250 m, 21.I.1965, MacKee 12034 (WELTU); Col d’Amieu, 350 m, 17.III.1965, MacKee 12272 Hypanthium (BISH, CANB, K[K001000422], MO, NOU[NOU029716], (1.5-)2-4.5 × 2-3 mm, campanulate or rarely obconic, sur- P[P00462985, P02088629], WELTU); Col d’Amieu, Route du face smooth, sparsely to moderately sericeous; ovary apex Col Toma, 400 m, 31.III.1965, MacKee 12342 (BISH, CAN- sparsely hairy. B[CANB00593372], K[K001000420], P[P00462986], WEL- TU); Mont Nakada pente Sud, 600 m, XI.1979, MacKee 37618 (MO[MO-6751048], NOU[NOU029714], P[P02089879]); Calyx N.C. Mont Yora (Pente Sud), 700 m, 4.XII.1980, MacKee 38405 Lobes 5, 1.5-2.0 mm, broadly rounded, glabrous above, gla- (NOU[NOU029560], MO[MO-6751050], P[P02089962, brous to sparsely sericeous below, green. P02089963, P05094544]); Mont Rembai, 700 m, 31.V.1990, MacKee 44919 (NOU[NOU029706]); Col d’Amieu, versant Canala, 350 m, 9.XII.1992, MacKee 46043 (NOU[NOU029557], Petals P[P02089842]); Forested slopes near Mt Rembai, south of Col 3.5-4.5 × 3.5-4 mm, ciliate but otherwise glabrous. d’Amieu, c. 500 m, 7.XI.1982, McPherson 5089 (BRI, MO[- MO-3218924], NOU[NOU029704], P[P00462976]); Mt Rembai Filaments region, above Col d’Amieu, c. 575 m, 7.VI.1983, McPherson 5734 3-5 mm; anther sacs 0.3-0.4 mm. (MO[MO-3211387], NOU[NOU029703], P[P00462977], PTNB, WELTU); Mt Rembai, N of Col d’Amieu, in the forest reserve SW of Canala-La Foa Rd, 650 m, 26.VII.1978, Phillips & Schmid Style 3205 (NOU[NOU53251], P[P00462978]); Col d’Amieu, Forêt de 4.5-6 mm, sparsely sericeous. Rembai, 500 m, 12.XI.1980, Suprin 876 (NOU[NOU029709]); Région de Pouembout, au n. de “Forêt-Plate”, 26.III.1981, Suprin Fruits 1087 (BISH fragment, NOU[NOU029708]); Col d’Amieu, horaire de la Foa, c. 350 m, V.1973, Veillon 2894 (NOU[NOU029713], 8-12 × 6.5-12 mm, subglobular to globular, base rounded, P[P00462979, P00462980, P00462981]; Col d’Amieu, Forêt Per- maturing yellow-green to reddish to reddish-black; seeds san, vers 400 m, 15.IV.1993, Veillon 7654 (NOU[NOU029707]). 4-12.

168 ADANSONIA, sér. 3 • 2020 • 42 (7) Revision of New Caledonian Gossia N. Snow & Guymer

165°E 166° N

Mt. Panié Ouvéa (1629 m) Island 21°S 21°S Koné

Mt. Humboldt Gossia pancheri (Brongn. & Gris) N. Snow (1618 m) Gossia ramiflora N. Snow, sp. nov.

22° Gossia virotii (Guillaumin) N. Snow 22°

100 km Nouméa

164°E 165° 166° 167°

Fig. 20. — Distribution maps of Gossia pancheri (Brongn. & Gris) N. Snow, G. ramifloraN. Snow, sp. nov. and G. virotii (Guillaumin) N. Snow.

Remarks the blades of the former are slightly bullate in some areas. These The species is easily diagnosed by the inflorescences on naked specimens also occur over sedimentary volcanics, in contrast to branches and elliptic, acute leaves with undulate margins. The the more typical substrates of volcanics and schists; the latter bark of branchlets often breaks into small rectangular flakes. specimen also is south of the known range of G. ramiflora sp. nov. Gossia ramiflora sp. nov. somewhat resembles G. clusioides, but the latter has a flat leaf margin, lowermost inflorescences axillary, and petals that are tomentose below. The species also resembles Gossia vieillardii (Brongn. & Gris) N. Snow G. vieillardii, but leaves of the latter mostly have flat margins, (Figs 19; 22; 23) are a lighter shade of green, and the fruit often tapers at the base. Specimens from Pic Noir north of Farino have shorter peti- Austrobaileya 8 (2): 181 (2010). — Eugenia vieillardi Brong. & Gris, Bulletin de la Société botanique de France 12: 180 (1865). — Austromyrtus oles, more rounded leaf bases, somewhat shorter inflorescences, vieillardii (Brongn. & Gris) Burret, Notizblatt des Botanischen Gartens und and slightly smaller flowers than average. The collection from Museums zu Berlin-Dahlem 15: 503 (1941). — Typus: New Caledonia. Pouembout is a shrub with more diminutive leaves. The margins Grande Terre, South Prov., Montagnes de Balade, 21°56’36.9960’’S, of the leaves of Munzinger 7207 are atypical in being sinuous 166°17’35.9880’’E, Vieillard 484 (lecto-, P[P00463006], designated only distally. by Snow (Snow & Veldkamp 2010: 181); isolecto-, G[G00340962], P[P00463007 specimen on left, P00463008]; see comments below). Two specimens previously identified asG. ramiflora sp. nov. may represent an undescribed species. They include Munzinger Myrtus prolixa Baker f., The Journal of the Linnean Society, Botany 7207 & Cantrill (MPU395687, P01031545) and Munzinger et al. 45: 311 (1921). — Austromyrtus prolixa (Baker f.) Burret, Notiz- blatt des Botanischen Gartens und Museums zu Berlin-Dahlem 15: 7261 (CANB n.v., MO-6630977, MPU028536, P01044671). 505 (1941). — Typus: New Caledonia. Grande Terre, South They resemble G. ramiflora sp. nov., but their leaf blades are Prov., Kaori forest in sheltered valley, Compton 393 (holo-, larger than normal with margins barely if at all irregular, and BM[BM000581653], photo BISH).

ADANSONIA, sér. 3 • 2020 • 42 (7) 169 Snow N.

Myrtus aneityensis Guillaumin, Journal of the Arnold Arboretum 12: The Rivière Blanche, Forêt du Mois de Mai, 25.VI.1951,Baumann - 254 (1931). — Austromyrtus aneityensis (Guillaumin) Burret, Bodenheim 14254 (MO, P, Z); région Rivière Bleue, 200 m, 27-28. Notizblatt des Botanischen Gartens und Museums zu Berlin-Dahlem III.1968, Bernardi 12412 (MO, US, Z); Thy House platform, 200 m, 15: 506 (1941). — Gossia aneityensis (Guillaumin) N. Snow, 26.II.1980, Brinon 549 (NOU[NOU029564]); Thy, upper eastern Novon 15: 478 (2005). — Typus: New Caledonia. Grande Terre, track, 290 m, 27.IV.1980, Brinon 646, (NOU[NOU029563]); Thy, South Prov., S. J. Kajewski 810 (lecto- [designated by Snow 2005: quarry track, 200 m, 23.X.1982, Brinon 1401 (NOU[NOU029562]); 181], P[P0646116]). — A later worker incorrectly annotated the Plaine des Lacs, c. 245 m, Compton 393 (BM); Goro, entre Forêt lectotype in 2011 as a syntype. Nord et le route Inco, 20.III.2002, Dagostini & Rigault 457 (NOU[NOU029618]); Goro-Inco, relevé R8, 20.IV.2002, Dagostini Eugenia heckelii Pancher & Sebert., Notice sur les Bois de la Nouvelle et al. 471 (NOU[NOU029612]); Goro-nickel, 20.XI.2002, Dagostini Calédonie: 259 (1874). — Typus: New Caledonia. Ab. loco, Petit & Rigault 602 (NOU[NOU029610]); Montagne des Sources, 1000 m, 52 (lectotype here designated, P[P004629880]). — Another 3.XII.1992, Dawson WELTU16579 (NOU[NOU029591], WEL- collection, Fournie 52 (P00462997) has “type?” written in the TU); Col. d’Amieu, 7.XI.1982, Dawson WELTU16600 (WELTU); upper right-hand corner of a label, but a later worker added “(no)” Port Boisé, 18.XI.1982, Dawson WELTU16604 (WELTU); Prony, without further explanation. 25.XI.1974, Favier 5195 (NOU[NOU029601]); Prony, II.1914, Franc 1689 (L, P[P005094037, P005094042, P005094047], US); Representative herbarium specimen. — P06668679. Prony, II.1914, Franc 1689a (P[P05094043, P05094045]); vallée supériure de la Pouéta-Kouré, 200 m, 20.V.1951, Guillaumin & Etymology . — Named for the French surgeon and botanist Eugène Baumann-Bodenheim 13274 (P[P00463019], Z); Montagne des Vieillard (1816-1896), who spent many years in New Caledonia. Sources, 22.VIII.1951, Hürlimann 1799 (P[P00463020], Z); vallée latérale de lat Riv. Blanche sur la route aux “Électriques”, 165 m, Distribution, habitat and phenology. — Common across 25.VIII.1958, Hürlimann 3377 (Z); Ouenarou Forest Reserve, much of Grande Terre and some islands (Fig. 19); in maquis of Koyama 8185 Le Rat humid forests or sometimes along streams, mostly over schists, mi- 280 m, 26.II.1992, (P[P00463022]); Prony, 306 caschistes, graywackes or serpentines, (5-)150-1000 m. Flowering (P[P05094051]); Baie de Sud [Baie de Prony], entre Baie January through June; fruiting known year-round. N’Go et Touaorou, 12.VIII.1903, Le Rat 600 (P[P05094046]); Mt. Dzumac, IV.1905, Le Rat 2659 (P[P05094048]); Haute Rivière Specimens examined. — New Caledonia. North Prov., Silva Peala, Bleue, 200 m, 8.IV.1965, MacKee 12396 (NOU[NOU029598], ad occidentem Pouebo, 500-600 m, 6.IV.1968, Bernardi 12609 ( P, P[P00463023], WELTU); Route de Yaté – Les Dalmates, 150 m, US, Z); ibid. loco, 200 m, 27-28.III.1968, Bernardi 12412 (L, P); La 8.III.1969, MacKee 20272 (NOU[NOU029613]); Col d’Amieu: Guen, refuge Blaffart, 25.II.2013, Hequet 4055 (NOU, MO); Ouest Toili, 600 m, 22.IX.1970, MacKee 22546 (NOU[NOU029561], de la Tiebaghi, 9.XI.1980, Hoff 3089 (NOU[NOU029575]); Ouest P[P02089837]); Montagne des Sources, 23.X.1974, 800 m, MacKee de la Tiebaghi, 9.XI.1980, Hoff 3108 (NOU); Ouraï, IX.1876, Lécard 29417 (P[P02089841]); Forêt Cachée, 250 m, VII.1973, MacKee 45 (P[P05094057]); Forêt de Tendé (Haute Diahot), 500-600 m, 26947 (NOU[NOU029617]); Prony, Forêt Nord, 200 m, 6.V.1975, 12.IX.1967, MacKee 17547 (NOU[NOU029597], WELTU; Haut MacKee 30090 (NOU[NOU029616]); Forêt Faux Bon Secours, Diahot – Tendé, Expl. Frouin, 500 m, 31.III.1969, MacKee 20456 1.V.1978, MacKee 35083 (NOU[NOU029570]); Pente Sud du (NOU[NOU029567]); Pouebo, crête entre Mandjélia et Salandané, Mt Nakada, 600 m, 18.XI.1979, MacKee 37618 (P[P02089879]); 600 m, 11.X.1969, MacKee 20819 (NOU[NOU029626]); Haute vallée Forêt Faux Bon Secours, 300 m, 17.III.1980, MacKee 37905 (NOU, d’Houailou, 350 m, 14.XI.1969, MacKee 21184 (NOU[NOU029715], P[P02089885]); Forêt Faux Bon Secours, 300 m, 25.II.1981, MacKee P[P02089836]); Pouebo, crête entre Mandjélia et Salandané, 38772 (NOU[NOU029571], P[P02089840]); Rivière Bleue, 150 m, 15.XII.1969, MacKee 21303 (NOU[NOU029568]); Pouebo, 600- 1.IV.1981, MacKee 38890 (NOU[NOU029569]); Vallee de la Ni, 700 m, 12.IV.1972, MacKee 25216 (BISH, NOU[NOU029625]); 100 m, 3.XII.1981, MacKee 40035 (MO, P[P02089886]); Rivière Col d’Amos, Haute Mayavetch, 500 m, 10.VII.1972, MacKee 25688 des Lacs 5 km aval de la Chute, 200 m, 24.I.1987, MacKee 43426 (P[P00463024, P05094542, P05094543]); Col d’Amoss, Haute (NOU[NOU029604]); Ouénarou, 150 m, 29.VIII.1987, MacKee Mayavetch, 500 m, 24.III.1973, MacKee 26472 (P[P02089835]); 43668 (NOU[NOU029603]); ibid. loco, 150 m, 7.III.1988, MacKee Aoupinié, 27.VII.1974, MacKee 29030 (NOU[NOU029614], 43923 (BRI, MO, NOU[NOU029609], P[P02089874]); Pic du P[P02089834]); [Île] Balabio, Baie de Capitaine, 10-150 m, 17.IX.1974, Pin, 200 m, 27.IX.1988, MacKee 44106 (P); Thy River valley, c. 12 MacKee 29317 (NOU[NOU029599]; P[P02089839]); Pouébo, air-km NE of Nouméa, 200 m, 9.IV.1979, McPherson 1539 (BISH, Mandjélia, 26.IV.1976, MacKee 31172 (NOU[NOU029607], MO, NSW, NOU[NOU029620], PTBG); Thy River valley,c. 12 air- P[P02089880]); Haut Diahot, Paala, 400 m, 1.VI.1976, MacKee 31296 km NE of Nouméa, c. 400 m, 25.IV.1979, McPherson 1567 (BISH, (NOU[NOU029615]); Haut Diahot, Tendé, 600 m, 16.V.1981, NOU[NOU029592], P[P00463027]); Along road from Dumbéa River MacKee 39055 (MO, NOU[NOU029605], P[P02089876]); Poné- valley towards Mt Dzumac, c. 25 air-km N of Nouméa, 24.II.1980, rihouen, Mont Aoupinié, 700 m, 18.VIII.1981, MacKee 39468 McPherson 2472 (MO, NOU[NOU029565], P[P00463028]); Rivière (MO, NOU[NOU029558], [P02089881]); Houailou, Néaoua, Bleue reserve, c. 30 air-km NE of Nouméa, 30.VII.1980, McPherson 450 m, 26.VI.1982, MacKee 40496 (MO, NOU[NOU029596], 2894 (MO, NOU[NOU029619], NSW, P[P00463029], WELTU); P[P02089877, P02089878]); Vallée de Bua Hio, 22.XII.1966, Schmid Rivière Bleue reserve, c. 200 m, 25.III.1981, McPherson 3648 (MO, 1929 (NOU[NOU029573], P[P00462994]); Mt. Mandjélia from NOU[NOU029621], P[P00463030], WELTU); Thy River Valley, Ouéga, c. 650 m, 11.VIII.2003, Snow 9214 et al. (ASU, BISH, BRI, c. 12 air-km NE of Nouméa, 25.III.1981, McPherson 3685d (MO); CANB, MO, NOU[NOU000646], NY, P[P00459486], WELTU); Rivière des Pirogues, N of Nouméa-Yaté road, 16.I.1982, McPherson Along streambed below and W of Col d’Amos, 22.X.1959, Thorne 4553 (MO, NOU[NOU029593], WELTU); Forested slopes near Mt 28143 (P[P00462999]); Massif Ignambi, 600 m, 28.XI.1967, Veil- Rembai, south of Col d’Amieu, 600 m, 6.XI.1982, McPherson 5064 lon 1513 (NOU[NOU053250], P[P00463000], Z); Paagoumène, (MO, NOU[NOU029623], P[P05094540]); Bay of Prony, 19.XI.1982, face site de l’usine, 13.V.1981, Veillon 4479 (NOU[NOU029600], McPherson 5160 (MO, NOU[NOU029566], P[P00463030], PTBG) ; P[P00463003]); Île Balabio, extremité sud, Bweroro, 5 m, 30.IX.1997, Forested slopes above a tributary of the Rivière Ni, 950-1000 m, Veillon 8035 (BISH, NOU[NOU029602], P[P00463004]); Wa- 10.XI.2003, McPherson 19212 et al. (BISH, NOU[NOU004200], MO, gape, 1867, Vieillard s.n. (NSW). — South Prov., Prony, Balansa 90 P[P06668679]); Exploitation Forestière Guiraud à l’est de Bourail, Me (P[P05094056], Z[Z-000092713, Z-000092714, Z-000092718]); Adeo, piste en direction du Menazi, 600-700 m, 24.III.1988, Morat Mois de May, 300 m, 25.VI.1951, Baumann-Bodenheim 14281 (A, 8008 (P[P00462987]); Pic du Pin, 7.XI.2002, Munzinger 1745 (MO, Z); Pte Panèta – Kouré, 10.V.1951, Baumann-Bodenheim 13274 (P); NOU[NOU012263], P[P00354256]); Haute Dumbéa, 25.V.1967,

170 ADANSONIA, sér. 3 • 2020 • 42 (7) Revision of New Caledonian Gossia N. Snow & Guymer

D A

E F I

Fig. 21. — A-D, Gossia pancheri (Brongn. & Gris) N. Snow: A, flowering branch with details of abaxial leaf venation offset( on right); B, detail of flower prior to anthesis; C, longitudinal section of flower bud showing bilocular ovary with axile placentation;D , leaf and fruits; E-I, Gossia ramiflora N. Snow, sp. nov.: E, flowering branch;F , details of more or less rectangular flaking bark;G , flowers prior to anthesis; H, detail of flower bud;I , another view of leaf (with detail of sinuous margin) with young fruits. Vouchers: A-C, McPherson 3275 (WELTU); D, McPherson 5019 (NOU); E-H, MacKee 12272 (WELTU); I, Phillips 2105 (NOU). Scale bars: A, D, I, 2 cm; B, C, H, 3 mm; E, 2 mm; F, G, 5 mm.

Nothis 373 (NOU[NOU029576]); Wagap, au-dessus de “Chapeau”, Schmid 5195 (P[P05094539], WELTU); Secteur du col d’Amieu, Pancher s.n. (P[P05094053]); Süd-Bai, zwischen der Bai N’Go und 700 m, 7.IV.1981, Suprin 1122 (NOU[NOU029595]); Monts Touaourou, 1903, Rohrdorf 12 (Z); Rivière Bleue, 11.III.1966, Dzumac, 26.XI.1981, Suprin 1542 (P[P00462998]); La Rivière Schmid 1097 (NOU[NOU029594], P[P005094538]); Rivière Blanche, V.1973, Veillon 2873 (NOU[NOU029622, NOU029599], Bleue, IV.1970, Schmid 3086 (NOU[NOU029574], P[P05094541, P[P00463001, P00463002]). — Province unknown/Ab loco. Franc P00462995]); Route du Dzumac, 600 m, 24.IV.1974, Schmid s.n. (P[P05094044]); Lécard s.n. (P[P05094055]); Sébert 74 (Herb. 5000 (NOU[NOU029572], P[P00462996]); Prony, 25.XI.1974, Pancher) (P[P05094052)]).

ADANSONIA, sér. 3 • 2020 • 42 (7) 171 Snow N.

Description contrast to my suggestion in 2010, the specimen should be Shrubs or trees considered the lectotype (as still annotated). 2-8(-18) m. Gossia vieillardii shows considerable variation in its morphology, but no character differences warranted splitting Branchlets the species. For example, a few specimens have slightly Terete, surface smooth; internodes > 5 mm. bullate leaves (e.g., Mackee 21184 and 22546). The species is the only one observed thus far in New Caledonia with Leaves occasionally tri-locular ovaries (Baker 1921). The petioles Coriaceous, 2 per node; petioles 2.5-6.5 mm; blades (2.0-)3.5- of G. vieillardii are bright pink or magenta. The base of the 8.0(-9.5) × (1.3-)1.8-3.5(-5.2) cm, narrowly elliptic to mostly fruit on specimens is either rounded or noticeably tapered. elliptic or broadly elliptic, base cuneate (rarely rounded), sur- The petals are whitish and said to be sweetly scented (Baker face flat, margin flat, apex obtuse to acute, midnerve above 1921). The flowers of some specimens are relatively small flush or sulcate, secondary veins above flush, oil glands of and have a densely pubescent hypanthium (e.g., MacKee lower surface absent or sparse, indistinct. 25216), whereas others are glabrous. The species can resemble G. diversifolia, but the latter has 4-angled branchlets. Inflorescence Gossia pancheri is similar but has a longer and more densely 1.5-5.0 cm, pedicels (1-)6-15(-30) mm; of monads, 3-flow- hairy hypanthium. ered cymes, or racemes, terminal or axillary, extrafloral bracts absent or scale-like. Gossia virotii (Guillaumin) N. Snow Bracteoles (Fig. 20) 0.5-1.4 mm, narrowly triangular, sparsely sericeous. Austrobaileya 8 (2): 192 (2010). — Myrtus virotii Guillaumin, Hypanthium Mémoires du Muséum national d’Histoire naturelle, Paris, sér. B, Bot. 4: 33 (1953). — Typus: New Caledonia. Grande Terre, South (2.5-)5-5.5 × 3.5-6 mm, obconic, surface smooth, sparsely Prov., Vallée de la Rivière de Humboldt, environs de la Case Marc sericeous; ovary apex glabrous to sparsely hairy. (ancien campement pages), 130 m, 13.XI.1940, Virot 411 (holo-, P[P00602573]; iso-, A[00255457], NOU[NOU006235], NY, Calyx P[P00462850], Z). Lobes 5, 1.4-2.7 mm, broadly ovate or rounded, ciliate on Specimens examined. — New Caledonia. South Prov., Baumann- margins, glabrous above, glabrous to sparsely sericeous be- Bodenheim 8119, Col de Vulcain, 11.XI.1950, (P[P00459517], US, low, green. Z); Col de Vulcain, 900 m, 11.XI.1951, Baumann-Bodenheim 8184 (P[P00459519], US); Col de Vulcain, 11.XI.1951, Bauman 8209 Petals (A, P[P00459520], US); Vale des von Mts Humboldt, 3.XI.1924, Däniker 459 (Z[Z-000050488]; Vallée Koéalagoguamba, 9.X.1977, (4-)5-7 × 3-4 mm, glabrous above and below, ciliate on margins. MacKee 34057 (NOU[NOU028911], P[P00462841]); Hte vallée de la Tontouta, 22.X.1968, Veillon 1857 (NOU[NOU028913], Filaments P[P00462844]). 3-5 mm; anther sacs 0.4-0.6 mm. Representative herbarium specimen. — P00462850.

Style Etymology. — After Robert Virot (1915-2002), French botanist 4-5 mm, sparsely (or rarely densely) villous on lower ½-3. and collector of the holotype.

Distribution, ecology and phenology. —Southern Grande Fruits Terre in maquis over ultramafics Fig( . 20), to c. 130 m. Flower- (5-)7-10(-12) × 6.5-10 mm, subglobular to globular, base ing August through February; fruiting September through March. rounded or often prominently tapered, red to dark red; seeds 1-2. Description Remarks Shrubs Gossia vieillardii is the most widespread species of the genus To 1 m tall. in New Caledonia. As presently understood, it is the only species of Gossia of New Caledonia that is not strictly endemic, Branchlets occurring also in Vanuatu (Snow & Veldkamp 2010). Round, internodes mostly greater than 5 mm long, emerging I annotated a specimen as the lectotype at P in August of pubescent becoming glabrous and grayish. 2004 but suggested later (Snow & Veldkamp 2010: 181) that it could be considered the holotype. Digital access to Petioles specimens at P, unavailable in 2004 and 2010, now reveals 0.5-2 mm, emerging hairy becoming glabrous. twelve specimens under Vieillard 484, representing at least four genera (Gossia, Decasperum J.R. Forst. & G. Forst., Leaves Uromyrtus Burret and Archirhodomyrtus [Nied.] Burret). In Coriaceous, 2-4 per node.

172 ADANSONIA, sér. 3 • 2020 • 42 (7) Revision of New Caledonian Gossia N. Snow & Guymer

A C

Fig. 22. — Gossia vieillardii (Brongn. & Gris) N. Snow: A, branch and inflorescences;B , flower just prior to anthesis;C , leaves and young fruits; D, leaves and young fruits; E, young fruit. Vouchers: A, B, McPherson 1567 (WELTU); C, Veillon 1513 (NOU); D-E, (MacKee 40566 [P]). Scale bars: A, C, D, 2 cm; B, 3 mm; E, 4 mm.

Petioles Anthers Less than 2 mm long. Said to be ovate (none seen).

Blades Fruit 1.5-3.0 × 0.2-0.6 cm, broadly linear to very narrowly Unknown. obovate, base cuneate, margins slightly recurved, apex retuse, sparsely hairy becoming more or less glabrous below, Remarks midvein sulcate above throughout, venation inconspicuous The species is recognizable by its frequent occurrence of 3-4 but secondary veins slightly raised above, inconspicuous to narrow leaves per node with their retuse tips. The species not visible below. resembles Gossia conspicua comb. nov., but G. virotii differs by its thinner textured, narrower and less strongly recurved Inflorescences leaf blades. In addition, the leaves of G. virotii are aggregated Of ascending axillary monads, infrequent, pedicels less than less densely at the tips of branchlets compared to G. conspicua 5 mm. comb. nov. The leaves ofBaumann-Bodenheim 8184 are sug- gestive of Gossia alaternoides, given their greater width than Calyx other specimens of G. virotii. Recent checks (August 2019) Lobes 4, whitish pubescent above. indicate that one specimen still is annotated (incorrectly) as G. alaternoides (P[P00462841]) and another (incorrectly) Petals as G. conspicua comb. nov. (P[P00462844]). Said to be whitish (none seen). Additional descriptive data are needed for fertile material, but despite the inadequate description of flowers (and Stamen disk none yet for fruits), the species is easily distinguished from Pubescent. G. conspicua comb. nov.

ADANSONIA, sér. 3 • 2020 • 42 (7) 173 Snow N.

Fig. 23. — Gossia vieillardii (Brongn. & Gris) N. Snow: A, branchlets in flower; B, mature (left) and immature (right) fruit and adaxial leaf surfaces. Vouchers: ©Hervé Vandrot, with permission; from forêt du Massif du Oua-Tilou (http://endemia.nc).

Synonymization of Gossia byrnesii under Eugenia Identification of sterile specimens of Myrtaceae typi- reinwardtiana cally is challenging, given that the family is species-rich in Described from a single specimen from northeastern Aus- many of the places where it occurs (e.g., Holst et al. 2014; tralia (Snow et al. 2003), Gossia byrnesii N. Snow & Guymer Sobral & De Souza 2015; Sobral et al. 2015; Snow et al. here is reduced to synonymy under Eugenia reinwardtiana 2015; Landrum 2017), which unfortunately also makes (Blume) DC., given its 4-merous flowers and dibrachi- distinguishing between different genera in the field and ate trichomes, the latter being common in Eugenia but herbarium difficult. Flowering material also may be difficult unknown for Gossia. Fruiting material, absent from the to identify at the generic level given that flowers of two or holotype, would have clarified unambiguously its generic more genera can be remarkably similar (e.g., Gossia, Eugenia, placement, given the pronounced differences in seed num- Archirhodomyrtus). In such cases fruits and mature seeds ber and testal texture (seeds few and with a soft or leathery typically are necessary to determine generic placement and testa in Eugenia). to make specific identifications. This conundrum is most The taxonomy ofE. reinwardtiana itself is complex and pronounced in New Caledonia between species of Eugenia in need of further study, summarized recently by Veldkamp and Gossia. Whereas members of Eugenia typically have (2013). Moreover, Bernardini et al. (2014) used maximum 4-merous flowers, some species of Gossia do as well (Snow likelihood analysis of combined nuclear (ITS and ETS) and et al. 2003). However, the hardened and relatively small, plastid (ndhF, matK, and rpl16) on six accessions of E. rein- usually somewhat compressed and irregularly angled seeds wardtiana from across a broad geographical area (Philippines of Gossia are distinguished easily from the relatively large, south to Australia and east to Hawaii). The sampled specimens globular, and soft-textured seeds of Eugenia. If a specimen were found to be closely related to, or perhaps conspecific bears dibrachiate (branched once near the base) trichomes, with, Eugenia bryanii Kaneh., E. palumbis Merr., E. confusa then it is Eugenia. If the trichomes are of other types, then DC., and E. koolauensis O. Deg. (Bernardini et al. 2014). Gossia is the more likely genus among the 4-merous baccate Myrtaceae in New Caledonia. However, some species of Eugenia in New Caledonia have other types of trichomes DISCUSSION (but see for example Snow et al. 2016a) in addition to the dibrachiate type (see also Snow et al., in prep.). Gossia is one of eight New Caledonian genera of Myrtaceae The other baccate genera in New Caledonia that share the with baccate fruits. Syzygium Gaertn. (Dawson 1999) and hardened and relatively small seeds of Gossia include (by Eugenia L. (flowers mostly 4-merous) are the most diverse, decreasing levels of specific diversity): Uromyrtus (c. 8-10 but both have relatively large, globular embryos with a species in New Caledonia; a widespread genus still badly in membranous to leathery testa. In addition, fruits of Syz- need of revision), which has solitary and strongly pendulous gyium and Eugenia in New Caledonia typically have three flowers with basifixed (non-versatile) anthers and distinctly or fewer seeds. In New Caledonia, Syzygium is most easily prolonged connectives on the filaments;Kanakomyrtus N. distinguished from Eugenia in having mostly paniculate Snow, a dioecious genus of six species with lobed stigmas inflorescences Dawson( 1999). (Snow 2009); Archirhodomyrtus (Nied.) Burret, a genus of

174 ADANSONIA, sér. 3 • 2020 • 42 (7) Revision of New Caledonian Gossia N. Snow & Guymer

three species (in N.C.) with a peltate stigma and numerous indetermined across various herbaria (pers. obs.), and the seeds stacked horizontally like poker chips in 2-3 well-defined accumulation of large numbers of digital images, the descrip- locules (vs stigmas narrow and terete and seeds typically five tive and distributional information in this treatment may or fewer arranged irregularly on shield-shaped placenta in need substantial updating. The author requests that new, Gossia), Rhodomyrtus locellata (Guillaumin) Burret, with large vouchered data be sent via email, preferably with detailed whitish-pinkish flowers and acrodromous leaf venation (vs observations and digital images, which will be incorporated shorter, whitish-pinkish petals and brochidodromous vena- (with acknowledgments) into the third volume of Myrtaceae tion in Gossia); Rhodamnia andromedoides Guillaumin (with of the Flore de la Nouvelle-Calédonie. [uniquely for the genus] 5-merous flowers, parietal placentation, and acrodromous venation on leaves that bear a short but stiff mucro); the monotypic Myrtastrum rufopunctatum Acknowledgements (Panch. ex Brongn. & Gris) Burret, a widespread, densely Extra special thanks to Jérôme Munzinger (MPU) for checking branched shrub (0.5-3 m) with unisexual flowers (plants the identify of many specimens, producing the distribution evidently usually dioecious; Snow in prep.) and with densely maps, and his thorough review that considerably improved ferrugineous-punctate leaves only c. 5-12 mm long; and Xan- the paper. Thanks also to the National Geographic Society thomyrtus kanalaensis (Hoch.) N. Snow, with 4-merous flow- for support of fieldwork in New Caledonia in 2003, and to ers (in New Caledonia) and a leathery and wrinkled mature the School of Graduate and Continuing Studies at Pittsburg fruit that resembles a raisin (Snow & Veldkamp 2010: 179). State University for summer support. The Muséum nacional As currently understood, Gossia in New Caledonia now d’Histoire naturelle in Paris provided generous financial sup- includes twenty species and ten subspecies. Together with port; Odile Poncy (P), Pete Lowry (MO) and Pete Phillipson species from Australia (nineteen species [now excluding G. (MO) facilitated a five week visit to P in 2015, where Matheiu byrnesi]) and elsewhere in Melanesia (six species), the genus Donnat assisted annotating and resorting hundreds of speci- now includes forty-five species and ten subspecies (Snow et al. mens. Kanchi Gandhi (GH) advised on several nomenclatural 2003; Snow 2005, 2006; Snow & Wilson 2010; Snow & matters. Assistance in the field in 2003 was provided by John Veldkamp 2010). Apart from G. vieillardii, which also occurs Dawson, Barry Sneddon, and Bettony Williamson. Bobbi An- in Vanuatu, all taxa of New Caledonian Gossia are endemic. gell provided most of the illustrations; Krista Anandakuttan of Gossia vieillardii also is by far the most common member of Illustrate Science provided those for the subspecies of Gossia the genus on Grande Terre; its absence from the Loyalty Is- clusioides. Gordon McPherson (MO), Martin Callmander lands (between Grande Terre and Vanuatu) is puzzling, given (G) and James Byng (L) shared data, checked specimens, and that it has reached Balabio Island to the north of Grande responded to numerous queries. Merci beaucoup for allowing Terre. Members of Gossia in New Caledonia occur mostly use of their digital images to Dominique Fleurot, Jean-Louis in forests, but over a wide range of substrates, precipitation Ruiz and Hervé Vandrot, and those who continue to send regimes, and elevations. images of other genera of Myrtaceae. Thanks to the curators Several species of Gossia are quite common (>10 collec- at A, BISH, BM, BRI, CAEN, CANB, G, GH, K, L, MEL, tions), including G. alaternoides, G. conspicua comb. nov., MPU, MO, NOU, NSW, NY, P, PTBG, RSA, WELTU, and G. diversifolia, G. kaalaensis sp. nov., G. pancheri, G. rami- Z for access to specimens, sending loans, or answering ques- flora sp. nov. and G. vieillardii; all likely will be designated tions. Finally, thanks to an anonymous reviewer for providing as Least Concern following IUCN Red List Categories the contructive comments. (IUCN 2012). The first preliminary and partial phylogeny ofGossia , based on nuclear ribosomal and plastid spacer DNA sequence REFERENCES data, suggests the genus arose in Australia but radiated soon thereafter both in New Caledonia and Australia (McLay Baker E. G. 1921. — Plants from New Caledonia. Dicotyledons. et al. 2018). The eight taxa sampled for New Caledonia were Polypetalae. The Journal of the Linnaean Society, Botany 45: 264- 323. https://www.biodiversitylibrary.org/page/171160 monophyletic, although their data set did indicate equivo- Beentje H. 2010. — The Kew Plant Glossary. Royal Botanic cal support for the separation of Austromyrtus from Gossia Gardens, Kew, 184 p. (McLay et al. 2018), despite consistent differences in floral Bernardini B., Byng J. W. & Snow N. 2014. — Phylogeny, morphology between the two genera (Snow et al. 2003). biogeographic origins, and convergent evolution in Old World An interesting ecological aspect of Gossia is the ability of Eugenia (Myrtaceae), with a focus on African species. Botany. Boise, Idaho, United States, poster with published abstract. several species to hyperaccumulate manganese (Fernando Brongniart A. & Gris A. 1865. — Observations sur les Myrt- et al. 2009, 2013; McLay et al. 2018). This ability may have acées sarcocarpées de la Nouvelle-Calédonie, et sur le nouveau helped broaden it ecological range in New Caledonia, given genre Piliocalyx. Bulletin de la Société botanique de France 12: the widespread presence of ultramafic substrates. 174-187. https://doi.org/10.1080/00378941.1865.10825005 Priorities for future work should focus on obtaining more Brown R. W. 1956. — Composition of Scientific Words. 2nd ed. Smithsonian Institution Press, Washington, 882 p. collections of the relatively uncommon taxa, and more of Burret M. 1941. — Myrtaceen-Studien. Notizblatt Botanis- the subspecies of Gossia clusioides. Given the rapid pace of che Gartens Berlin-Dahlem 15: 479-550. https://doi.org/ collecting in recent years, the many specimens that remain 10.2307/3995084

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E. & Siegele R. 2013. — Multiple metal accumulation onomique FLORICAL et les caractéristiques de la flore vasculaire within a manganese-specific genus. American Journal of Botany indigène de la Nouvelle-Calédonie. Adansonia, sér. 3, 34 (2): 100: 690-700. https://doi.org/10.1071/SB18016 177-219. https://doi.org/10.5252/a2012n2a1 Gâteblé G., Barrabé L., McPherson G., Munzinger J., Myers N. 1988. — Threatened biotas: “hot spots” in tropical Snow N. & Swenson U. 2018. — One new endemic plant forests. Environmentalist 8: 187-208. https://doi.org/10.1007/ species on average per month in New Caledonia, including BF02240252 eight more new species from Île Art (Belep Islands), a major Schlechter R. 1908. — Beitrage zur Kenntnis der Flora von micro-hotspot in need of protection. Australian Systematic Neu-Kaledonien. Botanische Jarbücher fur Systematik, Pflanz- Botany 31: 448-480. https://doi.org/10.1071/SB18016 engeschicte, und Pflanzengeographie, Beibl. 92, 40 (3): 22-45. Guillaumin A. 1911. — Histoire des explorations botaniques Sebert H. 1874. — Notice sur les bois de la Nouvelle Calédonie. ex Nouvelle-Calédonie et aux Îles Loyalty. Annales du Musée Arthus Bertrand, Paris, viii + 275 p. https://doi.org/10.5962/ colonial de Marseille, sér. 2, 9: 83-290. bhl.title.69241 Guillaumin A. 1931. — Contribution to the flora of the New Snow N. 1997. — Application of the phylogenetic species con- Hebrides: Plants collected by S.F. Kajewski in 1928 and 1929. cept: a botanical monographic perspective. Austrobaileya 5: 1-8. Journal of the Arnold Arboretum 12: 221-264. https://www. https://www.jstor.org/stable/41729913 jstor.org/stable/43780599 Snow N. 2005. — Five new combinations of Gossia in the Myr- Guillaumin A. 1938. — Contributions à la Flore de la Nouvelle- tle Family (Myrtaceae) from Australasia. Novon 15: 477-478. Calédonie. LXVII. Plantes récoltées par M. et Mme Le Rat de Snow N. 2006. — New species of Gossia N. Snow & Guymer 1900 à 1910 (10e supplement). Mémoires du Muséum national and Rhodomyrtus (DC.) Hassk. (Myrtaceae) from Papua New d’histoire naturelle, sér. 2, Botanique, 10: 433-434. Guinea. Austrobaileya 7 (2): 325-340. https://www.jstor.org/ Guillaumin A. 1939. — Matériaux pour la flore de la Nouvelle- stable/41739039 Calédonie. LIII. Revision des Myrtacées à fruit charnu précédée Snow N. 2009. — Kanakomyrtus (Myrtaceae): A new endem- de quelques notes supplémentaires sur les Myrtacées à fruit sec. ic genus from New Caledonia with linear stigma lobes and Bulletin de la Société botanique de France 8: 626-653. https:// baccate fruits. Systematic Botany 34: 330-344. https://doi. doi.org/10.1080/00378941.1938.10837451 org/10.1600/036364409788606253 Guillaumin A. 1959. — Contributions à la Flore de la Nouvelle- Snow N. & Veldkamp J. F. 2010. — Miscellaneous taxonomic Calédonie. CXVI. Plantes recueilles par MacKee (suite) 4e liste and nomenclatural notes for Myrtaceae. 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Snow N., Callmander M. C. & Phillipson P. 2015. — Stud- Sobral M., Faria Jr. J. E. Q., Ibrahim M. U., Lucas E. J., Rigui- ies of Malagasy Eugenia – IV: Seventeen new endemic species, era D., Stadnik A. & Villaroel D. 2015. — Thirteen new one new combination, and three lectotypifications; with com- Myrtaceae from Bahia, Brazil. Phytotaxa 224: 201-231. https:// ments on emerging distributional, ecological and evolution- doi.org/10.11646/phytotaxa.224.3.1 ary patterns. PhytoKeys 49: 59-121. https://doi.org/10.3897/ Thiers B. 2019 (and updated continuously). — Index Herbariorum: phytokeys.49.9003 A global directory of public herbaria and associated staff. New York Snow N., Dawson J. W., Callmander M. W., Gandhi K. & Botanical Garden’s Virtual Herbarium. http://sweetgum.nybg.org/ih/ Munzinger J. 2016a. — New species, new combinations, and Turland N. J., Wiersema J. H., Barrie F. R., Greuter W., lectotypifications in New Caledonian Eugenia L. (Myrtaceae). Hawksworth D. L., Herendeen P. S., Knapp S., Kusber W.- Candollea 71: 67-81. https://doi.org/10.15553/c2016v711a9 F., Li D.-Z., Marhold K., May T.W., McNeill J., Monro Snow N., Munzinger J. & Callmander M. W. 2016b. — Novi- A.M., Prado J., Price M. & Smith G. F. 2018. — International tates neocaledonicae V: Eugenia plurinervia (Myrtaceae): a Criti- Code of Nomenclature for Algae, Fungi and Plants (Shenzen Code). cally Endangered new species with distinct leaves. Candollea 71: Koeltz Scientific Books, Königstein. 201-205. https://doi.org/10.15553/c2016v712a7 Veldkamp J. F. 2013. — Nomenclatural notes on Eugenia reinward- Sobral M. & de Souza M. A. D. 2015. — Thirteen new Amazonian tiana (Myrtaceae) and more or less associated names. Garden’s Myrtaceae. Phytotaxa 238: 201-229. https://doi.org/10.11646/ Bulletin Singapore 65: 117-134. http://www.biodiversitylibrary. phytotaxa.238.3.1 org/page/43642528

Submitted on 28 June 2018; accepted on 21 September 2019; published on 27 April 2020.

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